Dr.  Lilienthal,  3 

230  W.  25th  St.,  New  York. 


MEMCAL    ^S€1HI®©L 


ittp://www.archive.org/details/cellcK)ctrineitshOOtysorici 


rig.  /.     J^g.  i. 


Fig.S. 


rtg.i. 


JHg.S. 


Fig.  6. 


Fig.  7. 


Youngest  lajer.  Middle  layor.  \^^     Oldest  layer/ 

'^:^^;lX':^Z1:lt^r^'^'^r''^''''  *"  EpHhenaioem  no.,  section  throu^n  laye^Kpiu^elic 


Fig.  to. 


POKMATION  OP  PUS. 
ToiHustrate  the  change  In  germinal  matter  of  an  Knithelial  cell,  resulting  from  increased  nutrition,  showine  tha 
manner  m  wmcli  the  germinal  matter  oi  u  normal  cell,  ifsupplied  freely  with  pabulum,  may  ?iverlse  to  pus. 
/''-'/■  II.  Fig.  12.  pig_  ^S_ 


Young.      Fully  fonrod.         Young. 

TENDON.  CAKTILAGK 


t5|!^^?--SPr  „________ 

'^— ^  , -S5 

s^^&s^-   'r^sm-  f=i^^w^^m 

m^F^^ 

a,  Sareolein 

i)n.     1,,  <■ 

MUSCLK. 

Fig.  /«. 


ELASTIC  TISSUE. 
The  arrow  shows  the  direction  in  which  g«nulnal 
matt4ir  u  supposed  to  be  moviuj;. 


NKRVK. 


I>eTelopment  of  young,  dark -bordered  nerre  fibres,  at  an  eariy 
period,  sliowing  germinal  mattur  and  formed  material  of 
elementary  parts.     X  IdOO. 


Fig.  /7. 


■  centre  or  nucleolus 


Germinal  matterC  nucleus). 


iMfRBA. 
Pura  germinal  matter.    X5000. 


Coarse  of  pabulum 


Oldest  part  of  formed 
material. 


Younpest  part  of 
formed  materia!. 


PLATE  ILLUSTRATING  DR.  BEALE'S  VIEWS, 


THE  CELL  DOCTRINE 


ITS  HISTORY  AND  PRESENT  STATE. 


FOR    THE    USE    OF 


STUDENTS  IN  MEDICINE  AND  DENTISTRY. 


A  COPIOUS  BIBLIOGRAPHY  OF  THE  SUBJECT. 


JAMES    T^YSON,   M.D., 

Lecturer  on  Microscopy  in  tde  University  of  Pennsylvania,  and  on  Physiology 

IN  THE  Pennsylvania  College  of  Dental  Surgery;  Fellow  of  the 

College  of  Physicians  of  Philadelphia,  etc.  etc. 


A  COLORED  PLATE  AND  OTHER  ILLUSTRATIONS. 


PHILADELPHIA: 

LIKDSAY  &  BLAKISTOK 

18  70. 

R 


Entered  according  to  Act  of  Congress,  in  the  year  1870,  by 

LINDSAY  &  BLAKISTON, 

In  the  Clerk's  Office  of  the  District  Court  for  the  Eastern  District  of  Pennsylvania. 


r  r    r;:,;      ;^cA2T0N 'PEERS  OP, ^  "^  ;:     I 

:'    ^  r  '   'sKEtekfAlf  db    Ca.:  'PHlLA£)Ef  PHll/ 


TO 

THE     MEDICAL     CLASS 

OF 

THE    UNIVERSITY    OF    PENNSYLVANIA, 

THIS  LITTLE   VOLUME 

IS    RESPECTFULLY    INSCRIBED, 

BY 

THE   AUTHOK. 


1S422 


PREFACE. 


The  author  has  become  convinced,  by  several 
years'  intimate  intercourse  with  students  of  medi- 
cine, that  their  acquaintance  with  the  subjects  he 
has  endeavored  to  include  in  this  little  volume 
would  be  facilitated,  if  the  views,  which  are  now 
taught  and  scattered  throughout  the  often  expen- 
sive works  of  their  authors,  were  collected  in  a  con- 
venient form  for  study  and  reference.  Taking  it 
for  granted  that  a  knowledge  of  this  subject  is  of 
fundamental  importance  in  its  bearing  upon  the 
study  of  physiology  and  pathology,  and  stimulated 
by  the  frequent  inquiries  of  students  for  an  appro- 
priate source  of  information,  he  has  prepared  what 
he  now  submits  to  them. 

He  has  sought  to  obtain  a  continuous  history  of 
the  evolution  of  the  "  cell  doctrine  "  up  to  its  present 
state,  without  embarrassing  his  pages  with  a  large 
number  of  isolated  facts.     He  has  attempted,  how- 


VI  PREFACE. 

ever,  to  secure  a  completeness,  and  to  make  the 
w^ork  useful  to  physicians  and  others  engaged  in 
research,  by  careful  references,  and  the  addition  of 
a  bibliography,  which  he  has  sought  to  make  accu- 
rate and  extended.  Some  authors  may  have  been 
overlooked ;  such  the  writer  cordially  invites  to  send 
him  references  to  their  own  papers,  or  to  those  of 
others  they  believe  to  have  a  bearing  upon  the 
subject. 

332,  South  Fifteenth  Street. 
February,  1870. 


ILLUSTRATION^  S. 


Plate. — Illustrating  Dr.  Beale's  Views. 

Figs.  1  to  7.  Production  of  formed  material  from  germinal  matter 
in  epithelial  cells,  from  section  through  layer  of  epi- 
thelium covering  papillae  of  frog's  tongue. 

Figs.  7  to  11.  Formation  of  Pus. 

Fig.  11.  "  "     Tendon. 

Fig.  12.  "  "     Cartilage. 

Fig.  13.  "  "     Muscle. 

Fig.  14.  "  "     Elastic  Tissue. 

Fig.  15.  "  "     Nerve. 

Fig.  16.  Amoeba. 

Fig.  17.  Illustrating  Nutrition  of  Cell. 

Intercalated. 
Fig.  1.  Illustrating  Globular  Theory.     After  Virchow. 

Fig.  2.  Cellular  Tissue  from  the  Embryo  Sac  of  Chamaedorea 
Schiedcana  in  the  act  of  formation.     After  Schleiden. 

Fig.  3.  From  the  Point  of  a  Branchial  Cartilage  of  Rana  Escu- 
lenta.     After  Schwann. 

Figs.  4  to  12.  Formation  of  Nuclei  and  Cells  from  Molecules,  ac- 
cording to  Bennett. 

Fig.  12.  Diagram  of  the  Investment  Theory.     From  Virchow. 


Vlll  ILLUSTRATIONS. 

Fig.  13.  Formation  of  Pus  from  subcutaneous  connective  tissue. 
From  Virchow. 

Fig.  14.  Formation  of  Pus  from  interstitial  connective  tissue  of 
muscle.     From  Virchow. 

Fig.  15.  Development  of  Cancer  from  connective  tissue.     From 
Virchow. 

Fig.  16.  Connective  Tissue  Corpuscles  anastomosing  one  with  the 
other.     From  Virchow. 

Fig.  17.  Formation  of  Elastic  Tissue,  according  to  Virchow. 


THE  CELL  DOCTRmE. 


The  idea  that  animals  and  plants,  however  com- 
plex their  organization,  are  really  composed  of  a 
limited  variety  of  elementary  parts,  constantly  re- 
curring, was  appreciated  by  Aristotle,  while  it  ap- 
pears to  have  been  even  more  clearly  conceived  by 
the  acknowledged  father  of  medical  science,  Galen. 
Fallopius  of  Modena,  1523-1562,  to  whom  we  are 
indebted  for  our  knowledge  of  the  conceptions  of 
Galen  in  regard  to  these  "partes  similares"  or 
'^simplices,"  has  further  developed  the  subject  of 
general  anatomy  in  his  "  Lectiones  de  Partibus  Simi- 
laribus  Ilumani  Corporis."  But  these  "  partes 
similares"  of  Fallopius,  which  were  bone,  cartilage, 
fat,  flesh,  nerve,  ligament,  tendon,  membrane,  vein, 
artery,  nails,  hairs,  and  skin,  plainly  do  not  corres- 
pond with  the  "  elementary  parts"  or  "  cells"  of  the 
present  day.  They  were  ultimate  to  Fallopius,  as 
stated  by  Prof.  Huxley,  because  he  could  go  no 
further,  "  though  it  is,  of  course,  a  very  different 
matter  whether  we  are  stopped  by  the  imperfection 
of  our  instruments  of  analysis,  as  these  older  ob- 
servers were,  or  by  having  really  arrived  at  parts 
no  longer  analyzable."*     These  "partes  similares" 

*  The  Cell  Theory— a  Keview,  by  T.  H.  Huxley :  Br.  &  For- 
eign Med.-Chir.  Rev.  for  Oct.,  1853,  No.  xxiv. 

2 


14  THE    CELL    DOCTRINE. 

really  correspond  to  the  "tissues"  of  the  present 
day,  which  are  collections  of  elementary  parts.  The 
conceptions  of  these  older  writers  with  regard  to  the 
"vital  endowment"  or  "independent  vitality"  of 
their  similar  parts  or  tissues,  were  singularly  correct, 
and  correspond  almost  identically  with  those  held  by 
the  majority  of  physiologists  of  the  present  day. 

Further  than  this,  however,  the  anatomists  of  the 
period  of  Fallopius  could  not  go — not  because,  as 
we  now  well  know,  they  had  arrived  at  parts  no 
longer  analyzable,  but  because  of  their  imperfect 
means  of  analysis. 

It  is  probable  that  the  magnifying  properties  of 
lenses  were  known  to  the  Egyptians,  as  well  as  the 
Greeks  and  Romans,  over  2000  years  ago;  since 
a  table  of  refractive  powers  is  introduced  into  his 
"  Optics"  by  Ptolemy,  since  Aristophanes,  the  comic 
Athenian  poet  (B.C.  500),  speaks  of  "burning 
spheres  "  of  glass  as  sold  in  the  grocers'  shops  of 
Athens,  and  since  both  Pliny  and  Seneca  refer  to 
lenses  and  their  magnifying  properties;  while  lenses 
themselves  have  been  found  in  the  ruins  of  Nineveh, 
Herculaneum,  and  Pompeii.  But  it  is  quite  certain, 
also,  that  they  did  not  become  available  as  com- 
pound microscopes  until  about  1590,  when  the  Jan- 
sens,  father  and  son,  of  Holland,  are  said  to  have  in- 
vented the  compound  microscope.  Fontana,  in  1646, 
writes  that  he  had  invented  the  microscope  in  1618. 
Galileo,  as  early  as  1612,  is  said  to  have  sent  a  micro- 
scope to  King  Sigismund  of  Poland,  though  whether 
it  w^as  his  own  invention,  or  made  after  the  pattern 
of  another,  is  more  difficult  to  determine.     In  1685 


THE    CELL   DOCTRINE.  15 

Stelluti  published  a  description  of  the  parts  of  a  bee 
he  had  examined  with  the  microscope,  and  although 
George  Ilufnagle  is  said  to  have  published  in  Frank- 
fort, in  1592,  a  work  upon  insects,  illustrated  by  fifty 
copper  plates,  it  is  highly  probable  that  these,  as  well 
as  very  many  most  important  observations  made  after 
the  invention  of  the  compound  microscope,  were 
made  with  the  simple  instrument.* 

It  is  impossible  to  estimate  the  assistance  the 
microscope  has  been  to  us  in  opening  up  the  minute 
structure  of  animals  and  vegetables,  and  in  thus  af- 
fording a  reliable  basis  on  which  to  build  a  doctrine 
of  organization.  Prof.  Huxley  says,  "The  influence 
of  this  mighty  instrument  of  research  upon  biology, 
can  only  be  compared  to  that  of  the  galvanic  battery, 
in  the  hands  of  Davy,  upon  chemistry.  It  has  ena- 
bled proximate  analysis  to  be  uUimate.''1[  But  it  is 
more  than  this.  Since,  as  he  correctly  states,  it  has 
enabled  proximate  physical  analysis  to  become  ulti- 
mate, it  corresponds,  not  to  the  galvanic  battery 
alone,  but  to  all  the  appliances  made  use  of  in  ulti- 
mate chemical  analysis. 

The  time  prior  to  the  invention  of  the  compound 
microscope  may  be  considered  as  the  fast  period  in 
histology;  that  between  this  date  and  that  of  the  ob- 
servations of  Schleiden  and  Schwann  (1838),  inclu- 
sive, the  second  period ;  while  the  time  subsequent  to 

*  For  an  interesting  and  exhaustive  history  of  the  invention  of 
the  compound  microscope,  see  Das  Mikroskop,  Theorie,  Gebrauch, 
Geschichte  und  gegenwartigcr  Zustand  desselben.  Von  P.  Harting 
In  drei  Biinden.     Braunschweig :  1866.     Dritter  Band,  ss.  11-35. 

f  Huxley,  loc.  citat.,  p.  290. 


16  THE   CELL   DOCTRINE. 

these  observations  becomes  appropriately  the  third 
period.  Notwithstanding  the  imperfect  state  of  in- 
struments during  quite  two  hundred  years  from  this 
date,  a  flood  of  facts  was  added  to  our  knowledge  of 
the  minute  structure  of  living  things. 

Borellus,  of  Pisa,  seems  first  to  have  used  the  mi- 
croscope in  the  examination  of  the  higher  animal 
structures,  about  the  year  1656,  but  his  observations 
were  grossly  misinterpreted  in  his  attempt  to  adapt 
them  to  the  prevailing  idea  of  the  day,  that  diseases 
were  caused  by  animalculae  in  the  blood  and  tissues. 
As  a  result,  he  describes  pus  corpuscles  as  animalcules, 
and  even  says  he  has  seen  them  delivering  their  eggs. 

According  to  Boerhaave,  Swammerdam  had  recog- 
nized the  blood  corpuscle  in  1658. 

Malpighi,*  between  1661  and  1665,  had  witnessed 
the  circulation  of  the  blood,  and  had  published  ob- 
servations upon  the  minute  structure  of  the  lungs, 
which  he  had  even  compared  to  a  racemose  gland,t 
kidneys,  spleen,  liver,  and  membranes  of  the  brain, 
and  with  some  of  these  structures  his  name  has 
become  inseparably  associated.  In  1667,  Robert 
HookeJ  pointed  out  the  cellular  structure  of  plants, 
and  Malpighi§  further  elaborated  the  same  subject 
with  considerable  accuracy  in  his  "  Anatome  Planta- 


*  Malpighi,  Opera  Omnia.     Lond. :  1686. 

f  Fort,  Anatornie  et  Physiologie  du  Pournon,  considere  comme 
organe  de  Secretion.  Paris:  1867,  Preface:  or  a  notice  of  Dr. 
Fort's  book,  by  the  writer,  in  American  Journal  of  Medical 
Sciences,  October,  1869. 

J  Hooke,  Kob.,  Micrographia.     Lond.:  1667. 

§  Malpighi,  Anatome  Plantarum.     London:  1670. 


THE   CELL   DOCTRINE.  17 

rum,"  in  1670.  He  showed  that  the  walls  of  the 
"  cells,"  or  "  vesicles,"  were  separable,  that  they  could 
be  isolated;  and  gave  to  each  the  name  ^^utriculus,'* 
believing  also  the  "cell,"  or  "utriculus,"  to  be 
an  independent  entity.  The  latter  observer*  also 
recognized  the  blood  .corpuscle.  Leeuwenhoek,  in 
1687,t  described  these  corpuscles  with  considerable 
accuracy,  not  only  in  man,  but  also  in  the  lower 
animals.  He  also  demonstrated  the  capillaries,  ex- 
amined most  of  the  tissues,  and  made  the  discovery 
of  the  spermatozoids,  which  he  conceived  to  be  sper- 
matozoa or  sperm  animals,  and  of  different  sexes. 

l!^o  attempt,  however,  seems  to  have  been  intelli- 
gently made  at  building  up  the  tissues  by  an  ultimate 
physical  element,  to  correspond  with  the  "atom"  of 
the  inorganic  chemist,  prior  to  that  of  Haller.  He  re- 
solved the  solid  parts  of  animals  and  vegetables  into 
the  "^6/^6 "  (fibra),  and  an  '' organized  concreted'  To 
the  former  he  assigns  the  most  important  position, 
asserting  that  it  is  to  the  physiologist  what  the  line 
is  to  the  geometrician;  that  a  "fibre,"  in  general, 
may  be  considered  as  resembling  a  line  made  up  of 
points,  having  a  moderate  breadth,  or  rather  as  a 
slender  cylinder.f 

The  second  elementary  substance  of  the  human 
body  according  to  Haller,  the  "  organized  concrete," 
must  not  be  lost  sight  of,  as  appears  to  have  been 

*  Malpighi,  Opera  Posthuma.     London:  1697. 

f  Leeuwenhoek,  Opera  Omnia  seu  Arcana  Naturae  detecta. 
Tom.  ii,  p.  421.  Leyden :  1687.  Vel  Opera  Omnia,  &c.,  Lugd. 
Batav. :  1722. 

J  Haller,  Elementa  Physiologiae,  vol.  i,  lib.  i,  sect.  i.  Lausan. 
Helvet.:  1757. 


18  THE   CELL   DOCTRINE. 

the  case  with  many  eminent  authorities  who  have 
attempted  to  give  his  views.  This,  he  says,  is  a 
mere  gUie,  evasated  and  concreted,  not  within  the 
fibres,  but  in  the  spaces  betwixt  them,  in  illustration 
of  which  it  is  stated,  that  cartilages  seem  to  be 
scarcely  anything  else  besides  this  glue  concreted. 
But  these  views  of  Haller  were  clearly  not  based 
upon  microscopic  observation,  though  the  microscope 
had  been  for  some  time  in  use.  For  Haller  himself 
tells  us  that  the  fibre  is  invisible,  and  to  be  distin- 
guished only  by  the  "  mind's  eye," — invisibilis  est  ea 
abra,  sold  mentis  acie  distingidmiis*  No  allusion  to  the 
cell  beyond  the  imperfect  description  of  the  blood 
corpuscles  and  spermatozoids  appears  to  have  been 
made  by  Haller. 

Better  founded,  in  being  based  upon  observation, 
was  the  theory  of  Wolf,  and  it  contained  many  of 
the  elements  of  truth.  For  an  available  exposition 
of  these  views,  physiologists  are  much  indebted  to 
Prof.  Huxley,  who  in  the  able  review  already  cited, 
has  presented  them  as  agreeing  partially,  also,  with 

*  A  singular  discrepancy  exists  between  these  words  of  Haller 
and  those  found  in  both  the  Latin  and  English  editions  of  the 
"  elegant  compend  "  of  Haller's  works  printed  in  Edinburgh,  the 
former  in  1766,  and  the  latter  (an  edition  in  the  possession  of  the 
writer),  in  1779,  under  the  inspection  of  William  Cullen,  M.D. 
In  the  latter,  we  have  the  following:  <'The  solid  parts  of  animals 
and  vegetables  have  this  fabric  in  common,  that  their  elements,  or 
the  smallest  parts  we  can  see  by  the  finest  microscope,  are  either 
fibres  or  an  organized  concrete. "i 

1  First  Lines  of  Physiology.  By  the  celebrated  Baron  Albertus  Haller, 
M.D.  Translated  from  the  correct  Latin  edition,  and  printed  under  the 
inspection  of  William  Cullen,  M.D.     Edinburgh  :  1779. 


THE   CELL   DOCTRINE.  19 

his  own.  The  doctrine  of  Wolf,  as  given  by  Prof. 
Huxley,  is  as  follows:  "Every  organ  is  composed,  at 
first,  of  a  mass  of  clear  viscous,  nutritive  fluid,  which 
possesses  no  organization  of  any  kind,  but  is  at  moat 
composed  of  globules.  In  this  semi-fluid  mass,  cavi- 
ties (Blaschen,  Zellen)  are  now  developed;  these,  if 
they  remain  rounded  or  polygonal,  become  the  sub- 
sequent cells,  if  they  elongate,  the  vessels;  and  the 
processs  is  identically  the  same,  whether  it  is  exam- 
ined in  the  vegetating  point  of  a  plant,  or  in  the 
young  budding  organs  of  an  animal.  Both  cells  and 
plants  may  subsequently  be  thickened  by  deposits 
from  the  '  solidescible'  nutritive  fluid.  In  the  plant, 
the  cells  at  first  communicate,  but  subsequently  be- 
come separated  from  one  another;  in  the  animal, 
they  always  remain  in  communication.  In  each  case 
they  are  mere  cavities  and  not  independent  entities;  organi- 
zation is  not  affected  by  them,  but  they  are  the  visible  results 
of  the  action  of  the  organizing  power  inherent  in  the  living 
mass,  or  what  Wolf  calls  the  vis  essentialis.  For  him, 
however,  this  vis  essentialis  is  no  Archaeus,  but  -simply 
a  convenient  name  for  two  facts  which  he  takes  a 
great  deal  of  trouble  to  demonstrate :  the  first,  the 
existence  in  living  tissues  (before  any  passages  are 
developed  in  them),  of  currents  of  the  nutritious 
fluid  determined  to  particular  parts,  by  some  power 
which  is  independent  of  all  external  influence ;  and 
the  second,  the  peculiar  changes  of  form  and  com- 
position, which  take  place  in  the  same  manner."* 


*  Huxley,  loc.  citat.,  p.  293-4.     Wolf,  C.  F.,  Theoria  Genera- 
tionis,  1759.     Ed.  Nova,  1774. 


20  THE   CELL   DOCTRINE. 

Two  points  are  here  particularly  to  be  observed  as 
cardinal, — first,  the  non-independence  of  cells,  either 
anatomically  or  physiologically;  that  they  are  ef- 
fects, passive  results,  and  not  causes  of  a  vitalizing  or 
organizing  force;  second,  that  organization  takes 
place  from  the  "  differentiation  "  of  the  homogeneous 
living  mass  in  these  parts,  through  the  agency  of  the 
vis  essentialis  or  inherent  vital  force.  The  radical 
difference  between  these  principles  of  development 
and  those  generally  held  at  the  present  day,  will  be 
better  appreciated  when  these  latter  have  been 
worked  out.  An  acknowledged  error  may  also  be 
pointed  out, — the  probable  result  of  the  inferiority  of 
the  instruments  of  that  day — that  of  supposing  the 
cells  of  plants  to  communicate  when  in  their  youngest 
state. 

This  theory,  however,  full  as  it  was  of  original 
conception,  and  based  on  actual  observation,  seemed 
to  claim  little  attention,  and  would  have  been  still 
less  known  but  for  the  labors  of  Prof.  Huxley.  The 
"  fibre"  theory  of  Haller  was  still  further  expanded, 
and  that  fibres  were  the  groundwork  of  nearly  all 
the  tissues,  continued  the  prevailing  view,  until  the 
latter  part  of  the  eighteenth  century,  and  there  are 
few  of  the  older  physiologies  even  of  a  later  date, 
which  do  not  contain  an  account  of  it.  I^aturally, 
it  maintained  itself  longest  in  the  case  of  the  fibrous 
tissues,  since  the  appearances  of  these  tissues,  when 
examined  by  the  highest  powers,  are  those  of  struc- 
tures apparently  composed  of  fibres. 

The  reaction  which  took  place  at  the  date  referred 
to  against  the  "fibre"  theory,  culminated  in  the 


THE   CELL   DOCTRINE.  21 

"globular"  theory,  due  less  to  speculation  than  er- 
roneous methods  of  observation  and  imperfect  in- 
struments. Leeuwenhoek*  (1687)  early  announced 
the  "globular"  structure  of  the  primitive  tissues 
of  the  body,  but  the  "globule"  apparently  attracted 
little  notice  until  this  period  of  reaction  against  the 
"  fibre,"  when  it  claimed  the  attention  of  Prochaskaf 
(1779),  FontanaJ  (1787),  the  brothers  Wenzel§  (1812), 
Treviranusll  (1816),  Bauer1[  (1818  and  1823),  Heu- 
singer**  (1822),  MM.  Prevost  and  Dumas,tt  Milne- 
Edwardstt    (1823),   Hodgkin§§    (1829),  Baumgart- 


*  Leeuwenhoek,  op.  citat. 

f  Prochaska,  De  Structura  Nervorum.  Vind. :  1779.  Opera 
min.,  Pars  i. 

I  Fontana,  Sur  les  Poisons,  1787,  ii,  18;  Abhandlung  iiber  das 
Viperngift,  das  Amerikanische  Gift,  u.  s.  w.  Aus  dem  Italien. 
Berlin:  1787. 

^  Wenzel,  Joseph  and  Charles.  De  structura  cerebri.  Tubing. : 
1812. 

II  Treviranus,  Verraischte  Schriften,  Anatom.  und  Physiolog. 
Inhalts.  Bd.  i.     Gottingen:  1816. 

^  Bauer,  Philosoph.  Transac.  for  1818,  and  Sir  E.  Home's  Lec- 
tures on  Comparative  Anatomy.  Vol.  iii,  Lect.  iii.    London  :  1823. 

**  Heusinger,  System der Histologic.    Thl.  i, Eisenach:  1822-4. 

If  MM.  Prevost  and  Dumas,  Bibliotheque  Universelle  dcs  Sci- 
ences et  Arts,  T.  xvii. 

ll  Milne-Edwards,  Memoire  sur  la  Structure  Elementaire  des 
Principaux  Tissues  Organiques  des  Aniraaux.  Paris:  1823.  Also, 
Kecherches  Microscopiques  sur  la  Structure  Intimo  des  Tissues  Or- 
ganiques des  Animaux,  in  Ann.  des  Sci.  Nat.,  Dec,  1826. 

g^  Hodgkin,  in  Grainger's  Elements  of  General  Anatomy.  Lon- 
don :  1829.  Also,  Hodgkin  and  Fisher's  translation  of  M.  Edwards 
"  Sur  les  Agens  Physiques."  London:  1832.  Hodgkin 's  Lectures 
on  the  Morbid  Anatomy  of  the  Serous  and  Mucous  Membranes. 
London:  1836,  p.  26.    Am.  Ed.    Philada. :  1838,  vol.  i,  pp.  17-18. 


22  THE    CELL   DOCTRINE. 

ner*  (1830),  F.  Arnoldf  (1836),  DatroclietJ  (1837), 
Kaspail§  (1839);  all  except  Hodgkin  admitting  in 
greater  or  less  degree  the  importance  of  the  globule 
as  an  ultimate  physical  element ;  while  it  is  evident, 
also,  that  there  was  much  confusion  in  the  use  of 
terms, — the  words  globule,  granule,  and  molecule,\\ 
being  often  indiscriminately  used,  and  the  word 
globule  sometimes  used  to  indicate  what  is  now  clearly 
recognized  as  the  "cell." 

Prochaska,T[  in  1779,  described  the  brain  as  made 
up  of  globules  eight  times  smaller  than  blood  glob- 

*  Baumgartner,  R.  H.,  Lehrbuch  der  Physiologie  mit  Nutz- 
anwendung  auf  die  arztliche  Praxis.    1853. 

f  Arnold,  F.,  Lehrbuch  der  Physiologie  des  Menschen.  Erst. 
Theil,  Zurich:  1836. 

X  Dutrochet,  M6nioires  pour  servir  a  I'Histoire  Anatomiqueet 
Physiologique  des  Vegetaux  et  des  Animaux.  T.  i,  ii,  Atlas. 
Paris:  1837. 

g  Easpail,  Systeme  de  Chimie  Organique.  2e  cd.,  T.  i,  ii. 
Brus. :  1839.     Avec  Atlas. 

II  The  German  authors  of  this  period  and  even  more  recent 
times  (Henle,  1841,  Virchow,  1858),  at  least  in  speaking  of  the 
development  of  histology,  seem  to  use  indiscriminately  the  terms 
granule  or  molecule,  and  globule,  whereas  they  are  morphologically 
something  distinct.  A  globule  is  usually  held  to  be  a  body, 
which,  under  the  microscope,  is  more  or  less  spherical  in  form, 
possessing  a  bright  centre,  and  dark  outline, — the  width  of  this 
outline  being  directly  as  the  difference  between  the  refracting 
power  of  the  globule  itself  and  that  of  the  menstruum  in  which  it 
floats.  Thus,  the  dark  outline  of  a  globule  of  oil  floating  in  water 
is  wider  than  that  of  the  same  globule  floating  in  glycerine.  A 
granule  or  molecule,  on  the  other  hand,  is  indeterminate  in  size  and 
shape,  and  appears  as  a  mere  dot  under  the  highest  powers  of  the 
microscope.  It  is  true  that  what  appears  as  a  granule  under  a  low 
power,  may  appear  as  a  globule  under  a  higher. 

^  Prochaska,  Opera  Minora,  Part  I,  p,  342. 


THE   CELL    DOCTRINE.  23 

ules.  In  the  year  1801,  the  philosophic  mind  of 
Bichat  elaborated  his  excellent  classification,  but 
he  seems  to  have  made  no  original  investigations  in 
minute  structure,  or  to  have  adopted  any  special 
theory  of  an  ultimate  physical  element.  The  bro- 
thers Joseph  and  Charles  Wenzel,*  in  1812,  de- 
scribed the  brain  as  composed  of  globules  of  small 
size.  Among  the  earliest  histologists  worthy  of 
mention,  is  Treviranus,f  whose  elements,  according 
to  Henle,  were  first,  a  homogeneous,  formless  matter; 
second,  fibres;  third,  globules  (kugelchen).  Mr. 
Bauer,J  quoted  as  a  most  experienced  microscopic 
observer  by  Sir  Everard  Home,  in  1818,  and  again 
in  1823,  described  the  ultimate  globules  of  the  brain 
and  of  muscular  fibre  as  of  the  size  of  a  globule 
of  blood  when  deprived  of  its  coloring  matter,  or 
about  2  0^0  0  ^^  ^^^  i^^^  i^  diameter.  The  fibre  was 
excluded  as  an  ultimate  element  of  organization 
by  Heusinger§  in  1822-4,  who  started  all  tissues 
from  the  globule,  still,  however,  retaining  the  form- 
less material  of  Haller  and  Treviranus.  Heusinger 
formed  the  fibre  by  the  linear  apposition  of  his 
globular  elementary  parts,  and  even  explained  how 
canals  and  vessels  were  formed  by  a  similar  ar- 
rangement of  vesicles  which  had  originated  from  the 
globules.  The  account  given  by  Henle||  of  the 
method  in  which  Heusinger  built  up  his  fibres  and 
vessels  is  interesting,  and  is  worth  translating,  since 


*  Wenzel,  op.  citat.,  p.  24.  f  Treviranus,  op.  citat. 

X  Bauer,  op.  citat.  §  Heusinger,  op.  citat.,  p.  112. 

II  Henle,  Allgemeine  Anatomie.     Leipzig :   1841,  p.  128. 


24  THE    CELL   DOCTRINE. 

there  is  in  these  views  an  approximation  to  the  truth. 
"As  the  result  of  an  equal  contest  between  contrac- 
tion and  expansion,  there  arises  the  globule,  of  which 
all  organisms,  all  organic  parts,  are  originally  com- 
posed. By  a  stronger  exercise  (Spannung,  tension) 
of  power,  there  originates  from  the  often  mere  homo- 
geneous globule,  the  vesicle.  Where  in  an  organism 
globules  and  ^formless  mass  are  present,  the  globules 
arrange  themselves  according  to  chemical  (?)  laws 
and  form  fibres.  Where  vesicles  arrange  themselves, 
there  arise  canals  and  vessels.'^  In  the  latter  sen- 
tence one  cannot  fail  to  note  a  close  approximation 
to  the  truth,  though  the  facts  upon  which  the  theory 
was  based  are  partly  false  and  partly  misinterpreted. 
But  the  observations  and  writings  of  Milne  Ed- 
wards* may  be  looked  upon  as  having  given,  more 
than  any  other  author,  position  and  popularity  to  the 
"  globular  theory."  lie  examined  all  the  principal 
tissues,  and  announced  that  the  fibres  of  the  then  so- 
called  cellular  (fibrous)  tissues,  membranes  composed 
of  these  fibres,  muscle  and  nerve,  were  composed 
of  globules  of  about  the  same  size,  from  gJ^^  to 
■rsuo  of  an  inch  in  diameter;  whence  he  concluded 
that  these  spherical  corpuscles,  by  their  aggregation, 
constituted  all  organic  textures,  vegetable  or  animal, 
and  whatsoever  their  properties  or  functions.  There 
is  little  doubt  but  that  many  of  these  so-called  glob- 
ules described  by  Edwards  were  really  cells,  seen 
with  indifi:erent  instruments,  and  further  distorted  by 
the  glare  of  direct  sunlight. 

*  Edwards,  loc.  citat. 


THE   CELL   DOCTRINE.  25 

Similar,  as  regards  the  element  of  organization, 
were  the  views  of  Baumgartner*  and  Arnold,!  who 
built  up  the  cell  wall  by  the  apposition  of  globules 
(to  which  the  term  granules  would  now  perhaps  be 
appliedj),  so  as  to  constitute  a  membrane  within 
which  other  globules  (granules)  remained  to  consti- 
tute contents. 

Fio.  1. 


Fig.  1.   Illustrating  the  globular  theory. 

A,  Fibre,  composed  of  elementary  granules  (molecular  granules),  drawn 
up  in  a  line.  B,  Cell,  with  spherically  arranged  granules.  (After  Vir- 
chow,  slightly  modified.) 

The  error  of  Edwards  seems  to  have  been  clearly 
pointed  out  by  Dr.  Hodgkin,§  though  much  impor- 
tance was  still  attached  to  the  globule  as  an  element 
of  organization  (but  perhaps  from  this  time  forward, 
more  in  the  stricter  sense  of  the  term  granule),  which 
has  continued,  in  this  latter  sense,  to  the  present  day. 

From  the  foregoing  facts,  it  is  evident  that  for  some 
time  prior  to  the  year  1838,  the  cell  had  come  to  be 
quite  universally  recognized  as  a  constantly  recur- 
ring element  in  vegetable  and  animal  tissues,  though 
as  yet  little  importance  had  been  attached  to  it  as  an 
element  of  organization,  nor  had  its  characters  been 

*  Baumgartner,  loc.  citat  ;  also,  Virchow,  Cellular  Pathology, 
Am.  Ed.  of  Chance's  Translation.    Philada. :   1863,  p.  53. 

j-  Arnold,  loc.  citat. ;  also,  Virchow,  Cellular  Pathology,  Am. 
Ed.  of  Chance's  Translation.     Philada. :  1863,  p.  53. 

I  See  note  to  p.  22.  g  Hodgkin,  loc.  citat. 

3 


26  THE   CELL   DOCTRINE. 

clearly  determined.  As  stages  in  its  growing  impor- 
tance may  be  mentioned,  the  demonstration  of  the 
cellular  structure  of  plants  by  Robert  Hooke  in  1667, 
the  further  elaboration  of  this  subject  by  Malpighi, 
and  his  statement  that  each  "utriculus"  was  an  in- 
dependent entity,  the  description  of  Heusinger,  in 
1822,  of  the  mode  of  formation  of  vessels  by  the  appo- 
sition of  vesicles,  already  referred  to,  and  the  announce- 
ment, though  erroneous,  of  Dbllinger,  in  1828,  that 
the  body  is  built  up  of  blood  corpuscles  which  move 
in  wall-less  (wandlos)  channels  in  the  tissues. 

A  most  important  contribution  to  the  anatomy 
of  the  cell  was  now  made,  in  the  discovery  of  the 
"  nucleus,"  by  Dr.  Eobert  Brown,  of  Edinburgh  ; 
whose  paper,  "  Organs  and  Mode  of  Fecundation  in 
Orchid^se  and  Asclepiadese,"  appeared  in  the  Trans- 
actions of  the  Linnean  Society  of  London,  in  1833. 
He  failed,  however,  to  appreciate  its  importance, 
though  its  discovery  was  another  fact  added  to  those 
necessary  to  complete  the  data  on  which  has  been 
founded  the  so-called  "  cell  theory." 

Singularly  near  the  truth  did  Easpail*  approach, 
in  1837,  when  he  tells  us  that  in  the  condition  of 
development  there  are  vesicles  or  cells,  endowed 
with  life  and  the  property,  almost  unlimited,  of  pro- 
ducing out  of  themselves  other  cells  of  the  same 
structure  and  similar  endowments,  of  spherical  form, 
and  capable  of  taking  up  oxygen  when  exposed  to 
the  atmosphere;  that  the  cell  membrane  in  its  fresh 
state  is  structureless.     Yet  he  considers  the  organic 

*  Kaspail,  op.  citat. 


THE    CELL    DOCTRINE.  27 

cell  as  made  up  of  granules  or  atoms,  spirally  ar- 
ranged about  an  ideal  axis,  comparing  the  cell  with 
the  crystal  rather  than  the  ultimate  element  or  atom 
of  which  the  crystal  is  made  up,  and  speaks  of  or- 
ganization as  crystallization  in  vesicles  {crystalliza- 
tion vesiculaire). 

Similar  was  the  view  of  Dutrochet,*  who  divided 
the  component  parts  of  the  body  into  solids  smd  fluid. 
The  solids  were  formed  by  the  aggregation  of  cells  of 
a  certain  degree  of  firmness;  the  liquids,  as  the  blood, 
are  also  made  up  of  cells/ which,  however,  float  freely 
among  each  other,  and  there  are  also  tissues  in  which 
the  cells  are  so  feebly  united,  that  one  can  scarcely 
tell  in  what  class  to  place  them.  The  contents  of  the 
cell  may  be  more  or  less  solid,  but  the  highest  degree 
of  vitality  is  only  compatible  with  liquid  cell  contents. 
Muscular  fibres,  and  the  remaining  animal  fibres,  are  cells 
much  elongated.  And  he  considers  the  same  general  plan 
to  prevail  in  the  animal  and  vegetable.  The  approach  of 
both  of  these  observers  to  the  truth  is  striking. 
Both,  however,  either  failed  to  detect  the  nucleus  or 
to  attach  any  importance  to  it.  They  failed  also  to 
lay  down  a  law  of  organic  development.  Hence  their 
views  were  soon  forgotten. 

Since  the  discover}^  of  the  nucleus,  by  Dr.  Robert 
Brown,  in  the  vegetable  cell,  it  had  been  recognized 
by  many  observers  in  various  pathological,  as  well  as 
healthy  animal  cells,  and  in  the  germ  cell  or  ovule 
of  birds,  as  early  as  in  1825,  by  Purkinje;    while 


*  Dutrochet,  op.  citat. 


28  THE   CELL   DOCTRINE. 

Purkinje,*  Yalentin,t  and  TurpiuJ  had  actually 
called  attention  to  the  relations  of  the  animal  and 
vegetable  cell  to  each  other. 

The  preexistence  of  the  nucleus,  and  the  gradual 
development  of  the  cell  about  it,  Valentin-  had  at- 
tempted to  demonstrate  in  the  case  of  pigment  cells, 
C.  H.  Schultz  in  the  blood  corpuscle,  Rudolph 
Wagner  in  the  egg,  and  Henle  in  epithelium,  all 
before  the  work  of  Schleiden  had  appeared.  Valen- 
tin, too,  had  said,  when  describing  the  nucleus  of 
epidermic  cells,  that  they  reminded  him  of  the  nu- 
cleus of  the  cells  of  vegetable  tissues.§  ISTot  only 
this,  but  Armand  de  Quatrefages||  and  DuraortierTf 
had  actually  observed  the  origin  of  young  cells  from 
the  full  grown,  in  the  embryo  of  the  freshwater  snail, 
while  Valentin  had  furnished  examples  of  the  devel- 
opment of  fibres  out  of  cells  in  muscular  fibre,  and 
in  the  substance  of  the  crystalline  lens.  In  fact,  as 
stated  by  Dr.  Waldo  J.  Burnett,  in  his  admirable 
paper,**  Valentin  "  perceived  the  true  physiological 

*  Purkinje,  in  Kaschkow,  Meletemata  Circa  Mammalium  Den- 
tium  Evolutionem.     Diss.  Inaug.,  Wratis. :  1835,  p.  12. 

f  Valentin,  Ueber  den  Verlauf  und  die  Enden  der  Nerven,  aus 
den  Nov.  Act.  Nat.  Curios.,  vol.  xvii:  besonders  Abgedruckt. 
Bonn:   1836. 

J  Turpin,  Ann.  d.  Sci.  Nat.,  2.  ser.  vii,  207. 

g  Valentin,  Nov.  Act.,  N".  C,  xvii,  pt.  I,  p.  96. 

II  Quatrefages,  Annales  des  Sci.  Nat.,  2  ser.  ii,  p.  114. 

][  Dumortier,  Annales  des  Sci.  Nat.,  2  ser.  vii,  p.  129. 

**  Burnett,  W.  J.  The  Cell ;  its  Physiology,  Pathology,  and 
Philosophy,  as  deduced  from  original  investigations.  To  which 
is  added  its  history  and  criticism.  A  prize  essay,  read  before  the 
American  Medical  Association,  and  published  in  vol.  vi  of  its 
Transactions.     Philadelphia:  1853. 


THE   CELL   DOCTRINE.  29 

relations  of  cells  as  far  as  he  well  could  without  ap- 
prehending the  grand  fact  that  the  nucleated  cell  is 
the  fundamental  expression  of  organic  forms." 

SCHLEIDEN   AND   SCHWANN. 

It  was  reserved  for  Schwann  to  accomplish  this 
master  stroke  in  observation  and  generalization, 
through  the  intermediate  results  of  Schleiden,  with- 
out whose  observations  on  vegetable  structures, 
the  true  position  of  the  cell  would  probably  have 
remained  undetected  for  some  time  longer.  Schlei- 
den, in  1838,  clearly  pointed  out  the  formation  of 
cells  in  vegetable  structures,  according  to  a  single 
and  uniform  method,  and  elaborated  the  theory  of 
development  of  which  the  cell  was  the  unit,  and 
which  Schwann  immediately''  extended  to  animal  tis- 
sues. 

A  formidable  obstacle  for  some  time  in  the  way  of 
a  law  of  development,  applicable  to  animal  and  veg- 
etable tissues,  was  the  opinion,  long  entertained,  that 
the  growth  of  animals  whose  tissues  are  furnished 
with  vessels  is  essentially  different  from  that  of 
plants;  an  independent  vitality  being  ascribed  to  the 
elementary  particles  of  vegetables  growing  without 
vessels.  So  firmly  was  this  believed,  that  the  ovum 
which  exhibited  undoubted  evidences  of  an  actual 
vitality  at  one  period  of  its  growth,  was  said  by  all 
physiologists  to  have  had  a  plant-like  growth.  This 
obstacle  was  removed  in  1837,  by  Henle,*  who  showed 

*  Henle,  Symbolae  ad  Anatomiam  vill.  intest.     Berol. :  1837. 

3* 


30  THE   CELL   DOCTRINE. 

that  an  actual  growth  of  the  elementary  parts  of  epi- 
thelium took  place  without  vessels. 

Taking  up  the  nucleus  as  discovered  by  Robert 
Brown,  Schleiden,*  in  reference  to  its  function,  ap- 
plies the  name  cytoblast  (xwtoc,  a  cell,  (SXaffvoq^  a 
bud  or  sprout),  or  "  cell  bud,"  and  in  a  careful  study 
of  its  anatomy,  discovers  that  "in  very  large  and 
beautifully  developed  cyloblasts,  there  is  observed 
a  small,  sharply  defined  body,  which,  judging  from 
the  shadow  which  it  casts,  appears  to  represent  a 
thick  ring,  or  thick-walled  hollow  globule."t  One, 
two,  three,  and  even  four  of  these  may  be  present. 
Without  further  present  comment  than  that  these 
characters,  as  given  by  Schleiden,  are  by  no  means 
constant,  it  is  plain  that  what  is  commonly  known 
as  the  nucleolus  is  here  intended,  to  the  discovery  of 
which  we  are  therefore  indebted  to  him,  though 
Valentin  also  claims  its  discovery  at  an  earlier 
period.!  ^1^  further  states  that  the  observations  he 
has  made  upon  all  plants,  lead  him  to  the  conclusion 
that  these  small  bodies  are  found  earlier  than  the 
cytoblasts. 

According  to  Schleiden,  when  starch  is  to  be  em- 

*  Schleiden,  Beitrage  zur  Phytogenesis,  Muller's  Archiv,  1838, 
p.  ii ;  Contributions  to  Phytogenesis,  Sydenham  Soc.  Transl.,  p. 
233. 

f  The  term  nucleolus  or  nucleus-corpuscle  (Kernkorporchen), 
seems  to  have  been  first  applied  by  Schwann.  (See  Introduction 
to  Schwann's  Kesearches,  Syd.  Society's  Translation.) 

X  Valentin,  "Outline  of  the  Development  of  Animal  Tissues," 
in  Wagner's  Elements  of  Physiology,  Translated  by  Dr.  Willis. 
London  :  1844,  p.  214.  Leipzig :  1839  ; — where  he  refers  to  Val- 
entin's Repertorium,  vol.  i,  p.  143. 


THE   CELL   DOCTRINE.  31 

ployed  in  new  formations,  it  becomes  dissolved  into 
sugar  or  gum,  which  are  convertible  into  one  another. 
The  sugar  appears  as  a  perfectly  transparent  fluid, 
not  rendered  turbid  by  alcohol,  and  receiving  from 
tincture  of  iodine  only  so  much  color  as  corresponds 
to  the  strength  of  the  solution.  The  gum  is  some- 
what yellowish,  more  consistent,  less  transparent, 
and  coagulated  into  granules  by  tincture  of  iodine, 
assuming  a  pale  yellow  color  which  is  permanent. 
In  further  progress  of  organization,  in  which  the 
gum  is  always  the  last  fluid,  a  quantity  of  exceed- 
ingly minute  granules  appears  in  it,  most  of  which, 
from  their  exceeding  minuteness,  appearing  as  black 
points.  Here  it  is  that  organization  takes  place, 
though  the  youngest  structures  are  composed  of 
another  distinct  homogeneous,  perfectly  transpa- 
rent substance — so  transparent  as  to  be  invisible 
when  not  surrounded  by  opaque  or  colored  bodies, 
— and  continuing  thus  after  pressure.  This  he  calls 
vegetable  gelatine,  and  considers  as  slight  modifica- 
tions, pectin,  the  basis  of  gum  tragacanth,  and  many 
of  the  substances  usually  enumerated  under  the  term 
vegetable  mucus.  It  is  this  gelatine  which  is  ulti- 
mately, through  the  agency  of  the  nucleus,  converted 
into  the  actual  cell-wall,  or  structures  which  consist 
of  it  in  a  thickened  state,  and  into  the  matter  of 
vegetable  fibre. 

In  this  homogeneous  blastema  or  cytohlastema  (said 
to  be  most  easily  studied  in  the  embryo  sac,  and  in  the 
extremity  of  the  pollen  tube),  are  very  soon  precipi- 
tated or  developed  mucous  or  protein  granules,  when 
the  solution  immediately  becomes  cloudy  and  more 


32 


THE   CELL   DOCTRINE. 


B      o 


or  less  opaque.  Single,  larger,  more  sharply  defined, 
granules  next  become  apparent,  A,  ^g.  2,  constituting 
the  nucleoli,  about  which  the  smaller 
granules  soon  collect  to  form  the 
nuclei  or  cytoblasts,  B.  These  then 
grow  considerably  in  the  free  state, 
C,  but  so  soon  as  they  have  at- 
tained their  full  size,  a  delicate, 
transparent  vesicle  rises  upon  their 
surface,  assuming  the  relation  of  the 
watch  crystal  to  a  watch,  D,  E. 
Thus  is  constituted  the  j^oung  cell. 
The  vesicle  gradually  expands  and 
becomes  more  consistent,  and  with 
the  exception  of  the  cytoblast, 
which  always  forms  a  portion  of  it, 
the  wall  now  consists  of  gelatine. 
The  entire  cell  then  increases  be- 
yond the  margin  of  the  cytoblast, 
and  quickly  becomes  so  large  that 
the  latter  at  last  merely  appears  as 
a  small  body  inclosed  on  one  of  the 
side  walls.  In  this  manner,  we  have 
exogenous  free  cell  formation.  With- 
in these  cells,  again,  as  well  as  in 
the  homogeneous  blastema  about 
them,  new  cytoblasts  arise,  grow, 
and  form  young  cells  which  grow  and  fill  up  the  mother 
cells,  and  finally  cause  them  to  disappear.  This  is 
endogenous  cell  formation.  According  to  Schleiden 
"the  entire  growth  of  the  plant  consists  only  of  a 
formation  of  cells  within  cells."     No  other  method 


P  E 

Fig,  2.  Cellular  Tis- 
sue, from  the  embryo 
sac  of  Chamaedorea 
Schiedeana,  in  the  act 
of  formation. 

A,  Formative  sub- 
stance, gum,  mucous 
granules,  nuclei  of  cy- 
toblasts (nucleoli). 

B,  Cytoblasts. 

C,  Single  and  free  cy- 
toblast more  highly 
magnified. 

D,  Cytoblast  with 
cell  forming  in  it. 

E,  Same  more  highly 
magnified. 

F,  Cytoblast  isolated 
after  destruction  of  cell. 

From  Schleiden  — 
Beitrage  zur  Phy togen- 


THE   CELL   DOCTRINE.  33 

of  formation  of  new  cells  seems  to  have  been  con- 
ceived by  him.  For  although  the  multiplication  of 
cells  by  fissiparous  division  of  previously  existing 
cells,  had  been  demonstrated  by  Mirbel,*  and  con- 
firmed by  Von  Mohl,t  before  the  investigations  of 
Sehleiden  had  been  made,  the  latter  author  considered 
the  apparent  growing  across  of  the  partition  cells  an 
illusion,  and  that  the  young  cells  escape  observation 
in  consequence  of  their  transparency,  until,  at  a  late 
stage,  their  line  of  contact  is  regarded  as  the  partition 
wall  of  the  parent  cell;  while  even  Schwann  states 
somewhat  hesitatingly  what  is  now  so  generally  ad- 
mitted.J  This  is  the  cell  theory  of  Sehleiden,  which 
has  also  been  properly  called  the  theory  of  free  cell 
formation,  since  it  involves  a  spontaneous  generation 
of  the  cell. 

The  merit  of  Schwann  consisted  in  applying  this 
theory  to  animal  tissues,  his  conclusions  being  based 
upon  the  study  of  the  formation  of  the  chorda  dor- 
salis,  and  cartilage,  and  a  comparison  of  their  cells 
with  those  of  vegetable  tissues.  Thus,  in  a  cyto- 
blastema,  either  structureless  or  minutely  granulous, 
"a  nucleolus  is  first  formed;  around  this  a  stratum 
of  substance  is  deposited,  usually  minutely  granulous, 
but  not  yet  sharply  defined  on  the  outside.  As  new 
molecules  are  constantly  being  deposited  in  this  stra- 
tum between  those  already  present,  and  as  this  takes 
place  within  a  precise  distance  of  the  nucleolus  only, 

*  Mirbel,  Recherches  sur  la  Marchantia,  1833. 
f  Von  Mohl,  Entwicklung  und  Bauder  Sporen  der  Kryptogam. 
Gew.,  Flora,  1833. 

X  Schwann,  op.  citat.     Introduction,  p.  4. 


34  THE   CELL   DOCTRINE. 

the  stratum  becomes  defined  externally,  and  a  cell 
nucleus  having  a  more  or  less  sharp  contour  is  formed. 
The  nucleus  grows  by  a  continuous  deposition  of 
new  molecules  between  those  already  existing,  that 
is  by  intussusception.  (See  Fig.  3,  e.)  If  this  go  on 
equally  throughoutthe  entire  thickness  of  the  stratum, 

Fio.  3. 


Fig.  3.  From  the  point  of  a^branchial  cartilage  of  Rana  esculenta. 
(From  Schwann.) 

the  nucleus  may  remain  solid;  but  if  it  go  on  more 
vigorously  in  the  external  part,  the  latter  will  become 
more  dense,  and  may  become  hardened  into  a  mem- 
brane, and  such  are  the  hollow  nuclei."* 

When  the  nucleus  has  reached  a  certain  stage  of 
development,  the  cell  is  formed  around  it.  The  fol- 
lowing is  the  process  by  which  this  takes  place : — 
"A  stratum  of  substance,  which  differs  from  the  cyto- 
blastema,  is  deposited  upon  the  exterior  of  the  nu- 
cleus. (See  Fig.  3,  d.)  In  the  first  instance,  this 
stratum  is  not  sharply  defined  externally,  but  be- 
comes so  in  consequence  of  the  progressive  deposition 
of  new  molecules.  The  stratum  is  more  or  less  thick, 
sometimes  homogeneous,  sometimes  granulous:  the 

*  Schwann,  op,  citat.,  p.  176. 


THE   CELL    DOCTRINE.  35 

latter  is  most  frequently  the  case  in  the  thick  strata 
which  occur  in  the  formation  of  the  majority  of  ani- 
mal cells.  We  cannot,  at  this  period,  distinguish  a 
cell  cavity  and  cell  wall.  The  deposition  of  new 
molecules  between  those  already  existing  proceeds, 
however,  and  is  so  effected  that  when  the  stratum  is 
thin,  the  entire  layer,  and  when  it  is  thick,  only  the 
external  portion,  becomes  gradually  consolidated  into 
a  membrane.  The  external  portion  of  the  layer  may 
become  consolidated  soon  after  it  is  defined  on  the 
outside ;  but,  generally  the  membrane  does  not  be- 
come perceptible  until  a  later  period,  when  it  is 
thicker  and  more  defined  internally;  many  cells, 
however,  do  not  exhibit  any  appearance  of  the  for- 
mation of  a  cell  membrane,  but  they  seem  to  be  solid, 
and  all  that  can  be  remarked  is  that  the  external  por- 
tion of  the  layer  is  somewhat  more  compact.* 

"Immediately  that  the  cell  membrane  has  become 
consolidated,  its  expansion  proceeds  as  the  result  of 
the  progressive  reception  of  new  molecules  between 
the  existing  ones;  that  is  to  say,  by  virtue  of  a  growth 
by  intussusception,  while  at  the  same  timie  it  becomes 
separated  from  the  cell  nucleus The  in- 
terspace between  the  cell  membrane  and  the  cell 
nucleus  is  at  the  same  time  filled  with  fluid,  and  this 
constitutes  the  cell  contents.  During  this  expansion 
the  nucleus  remains  attached  to  a  spot  on  the  internal 
surface  of  the  cell  membrane."  Though,  according  to 
Schwann,  in  animal  cells  the  nucleus  is  never  covered 
by  a  lamella  passing  over  its  inner  surface,  as  is  the 

*  Schwann,  op.  citat,  p.  176. 


36  THE    CELL   DOCTRINE. 

case  with  the  vegetable  cell  according  to  Schlei- 
den.  Thus  is  formed  the  animal  cell  according  to 
Schwann,  and  although  its  method  is  identical  with 
that  of  Schleiden,  both  as  to  endogenous  and  ex- 
ogenous cell  formation,  we  have  quoted  his  own 
paper,  because  he  is  plainly  fuller,  and  more  precise 
in  his  descriptions.  The  object  of  each  observer  was, 
however,  the  same  with  regard  to  the  tissues  studied; 
the  additional  object  of  Schwann  being  to  show  that 
all  organisms,  ivhelher  animal  or  vegetable,  are  formed  on 
a  common  principle,  a7id  that  this  principle  is  origin  from 
cells, — that  the  various  tissues  of  the  plant  and  animal, 
however  simple  or  complicated,  are  all  combinations 
of  these  cells,  modified  in  adaptation  to  the  special 
peculiarities  of  tissues. 

The  conception  of  Schleiden  was  truly  original, 
though  its  application  was  less  difficult  in  conse- 
quence of  the  simplicity  of  vegetable  tissues.  The 
conception  of  Schwann  was  easier,  in  being  the  re- 
flection of  that  of  Schleiden,  while  its  application  was 
more  difiicult,  in  consequence  of  the  great  diversity  of 
animal  tissues;  so  difiicult  that  he  acknowledged  that 
"there  are  some  exceptions,  or  at  least  difterences, 
which  are  as  yet  unexplained."  This  need  not  sur- 
prise us  when  we  recollect  that  one  of  the  ablest 
modern  exponents  of  the  cell  theory,  admits  the  diffi- 
culty of  its  application  to  some  of  the  so-called  higher 
tissues.*  Indeed,  the  careful  reader  of  Schwann's 
researches  cannot  but  be  surprised  at  the  accuracy 
of  the  observations  of  this  histologist,  nor  can  he 

*  Virchow,   Cellular   Pathology,  Chance's    Translation.  Am. 
Edit.,  Philada. :  1863,  p.  78. 


THE   CELL   DOCTRINE.  37 

fail  to  realize  how  comparatively  few  have  been  the 
changes  necessitated  in  his  descriptions,  or  the  method 
of  application  of  his  theory  to  the  formation  of  the  dif- 
ferent tissues.  Indeed,  the  portion  of  the  theory  of 
Schleiden  and  Schwann  which  does  not  accord  with 
the  latest  expression  of  the  cell  doctrine,  is  not  so 
much  that  which  pertains  to  the  formation  of  tissues 
from  existing  cells,  as  that  which  relates  to  the 
method  in  which  they  supposed  the  cells  to  origi- 
nate; which,  it  will  be  recollected,  was  by  a  species 
of  spontaneous  generaiioii  of  the  essential  parts  of  the 
cell,  in  a  homogeneous  cytoblastema. 

A  difterence  in  the  anatomy  of  the  cell  as  given  by 
Schwann,  and  physiologists  of  the  present  day  is 
seen  in  the  location  of  the  nucleus  by  the  latter,  who 
places  it  not  merely  eccentrically,  but  actually  "  sepa- 
rated from  the  surface  only  by  the  thickness  of 
the  assumed  cell-wall;"*  though  an  inspection  of 
Schwann's  drawings  would  not  convey  this  impres- 
sion. At  the  present  day,  the  situation  of  the  nu- 
cleus, though  usually  central,  is  known  to  be  not 
unvarying.  Again,  the  primary  and  absolutely  es- 
sential presence  of  the  nucleolus,  as  well  as  the  uni- 
versal presence  of  the  cell-wall,  may  be  considered 
characteristics  of  Schleiden  and  Schwann's  idea  of 
the  cell,  which  are  now  no  longer  insisted  upon. 

As  already  stated  (p.  33),  Schwann  would  seem 
to  have  admitted  also,  the  formation  of  cells  by 
division,  though  with  some  hesitation.  Thus  he 
writes  :t  "A  mode  of  formation  of  new  cells,  differ- 

*  Schwann,  op.  citat.,  p.  37,  a.  f. 
f  Schwann,  op.  citat.,  Introduction,  p.  4. 
4 


38  THE    CELL    DOCTRINE. 

eiit  from  the  above-described,  is  exhibited  in  the 
multiplication  of  cells  by  division  of  the  existing 
ones;  in  this  case,  partition-walls  grow  across  the  old 
cell,  if,  as  Schleiden  supposes,  this  be  not  an  illusion, 
inasmuch  as  the  young  cells  might  escape  observa- 
tion in  consequence  of  their  transparency,  and  at  a 
later  stage,  their  line  of  contact  would  be  regarded 
as  the  partition-wall  of  the  parent  cell." 

It  would  be  easy  to  point  out  other  defects  in  the 
theory  of  Schleiden  and  Schwann,  when  it  is  tested 
by  comparison  with  the  more  accurate  observation 
of  the  last  twenty-five  years,  none  of  w^hich  should 
be  permitted  to  detract  from  the  credit  which  at- 
taches to  the  originators  of  this  conception.  It  must 
not  be  forgotten,  that  it  is  no  less  true  of  science  than 
of  art,  that  great  and  important  truths  in  their  en- 
tirety, are  gradually'  developed,  and  that  no  single 
mind  is  capable  of  elaborating  them  from  their  in- 
cipiency  to  their  complete  expression.  And,  since 
many  clever  people  had  daily  noticed  the  rising  of 
steam  from  the  boiling  kettle  without  thinking  of 
utilizing  its  principle  of  expansion,  so  many  careful 
observers  had  time  and  again  witnessed  the  cellular 
or  vesicular  composition  of  plants,  and  yet  failed  to 
appreciate  the  importance  of  the  nucleated  cell,  and 
to  deduce  from  it  a  law  of  development  applicable  to 
all  organic  forms.  Again,  as  the  engine  of  Watt 
was  far  different  from  the  beautiful  and  powerful 
creation  of  the  mechanic  of  the  present  day,  so  the 
cell  theory,  as  developed  by  Schleiden  and  Schwann, 
has  been  further  evolved  by  later  histologists.  So 
that  we  may  truthfully  reiterate,  with  Prof  Huxley, 


THE    CELL    DOCTRINE.  39 

"  whatever  cavillers  may  say,  it  is  certain  that  histol- 
ogy before  1838,  and  histology  since  then,  are  two 
difterent  sciences — in  scope,  in  purpose,  and  in  dig- 
nity— and  the  eminent  men  to  whom  we  allude, 
may  safely  answer  all  detraction  by  a  proud  '  circum- 
spiee.'  "* 

According  to  these  observers,  then,  a  perfectly- 
formed  cell  would  be  defined  as  a  closed  vesicle,  with 
certain  contents,  among  which  were  essentially  a  nu- 
cleolus and  nucleus, 

HENLE,   1841. 

It  is  not  consistent  with  our  object  to  include  all 
of  the  numerous  observations  which  were  multiplied 
after  this  period,  incited  by  the  researches  of  Schlei- 
den  and  Schwann.  It  is  simply  to  point  out  the 
salient  features  of  those  results  which  point  towards, 
and  have  culminated  in  accepted  views.  It  has 
been  stated  that  previous  to  Schleiden's  researches, 
in  1838,  the  formation  of  cells  by  division  had  been 
asserted  as  one  mode  of  origin  of  cells,  that  Schlei- 
den  had  declared  this  an  error  of  observation,  and 
that  Schwann  had  hesitatingly,  if  at  all,  accepted  it 
as  a  rare  method  of  cell  formation. 

IIenle,t  who,  in  general,  adopted  the  view  of 
Schwann  as  to  the  jmmary  origin  of  cells,  though  he 
made  exception  to  its  universality  of  application, 
says  that  cells  multiply  in  three  ways : 

1.  By  budding  (durch  Sprossen),  as  in  certain  lower 
plants. 

*  Huxley,  op.  cit.,  p.  290. 

•}•  Henle,  Allgemeine  Anatomie.     Leipzig  :  1841,  p.  172  et  seq. 


40  THE    CELL   DOCTRINE. 

2.  By  endogenous  cell  development  (durch  endo- 
gene  Zeugung),  where  the  cell  contents  of  the  mo- 
ther cell  become  the  cj^toblastema  of  the  daughter 
cells,  as  originally  given  by  Schleiden  and  Schwann. 

3.  By  division  or  segmentation  (durch  Theilung), 
of  which  he  says,  however,  no  examples  are  found 
among  animals;  though  he  also  states  in  the  para- 
graph* immediately  following,  "We  would,  with 
Schwann,  consider  cell  formation  in  the  yolk,  by 
*  farrowing,'  an  analogous  process,  if  we  may  con- 
sider the  yolk  as  a  simple  cell."  He  then  proceeds 
to  describe  how,  by  a  "cording  in"t  of  the  surface, 
the  yolk  is  divided  into  two  equal  parts,  these  into 
four,  and  so  on  until  the  entire  yolk  becomes  a  mul- 
berry mass,  made  up  of  little  round  bodies.  This  seg- 
mentation of  the  ovum  already  observed  in  the  yolks 
of  frogs,  fish,  molluscs,  and  medusae,  Henle  says  at 
this  time  (1841),  has  perhaps  merely  escaped  noticej 
in  the  case  of  the  higher  animals,  as  plausibly  sus- 
pected by  Bergman, — a  suspicion  which  we  need 
scarcely  say  was  amply  confirmed  a  little  later.  But 
Henle  also  states,  in  the  same  connection,  that  certain 
cases  arise  in  which  perfect  cells  are  developed  in  a 
cytoblastema,  in  a  manner  w^hich  is  inexplicable, 
and  that  from  these  cells,  tissues  are  finally  devel- 
oped. §  Whence  the  undetermined  state  of  the  question 


*  Henle,  op.  cit.,  p.  176. 

I  Einschniiren,  to  cord  (a  box,  &c.),  to  bind  with  a  cord,  to 
string.     Orieb. 

X  Henle,  op.  cit.,  p.  177. 
§  Henle,  op.  cit.,  p.  177. 


THE    CELL    DOCTRINE.  41 

at  that  time  may  be  easily  inferred.*  J^or  is  mention 
here  made  by  Henle,  of  the  nucleus  of  the  cell,  as  the 
primary  seat  of  the  segmentation.  The  surface  of  the 
cell  is  said  to  be  "  corded  in,"  or  "furrowed,"  deeper 
and  deeper,  until  the  division  takes  place.  This  de- 
scription is  still  adhered  to  by  many  physiologists 
of  the  present  day,  who  consider  that  there  is  a  sim- 
ple disappearance  of  the  germinal  vesicle  or  nucleus 
of  the  ovum  after  fecundation,  rather  than  a  division 
of  it  into  two,  and  substitution  of  these  for  the  orig- 
inal one. 

*  While  endeavoring  to  trace  out  the  steps  by  which  the  present 
most  generally  accepted  views  with  regard  to  the  origin  of  cells,  were 
arrived  at,  it  must  not  be  forgotten  that  other  dissenting  views  were 
also  advanced,  though  tending  differently  from  those  incorporated 
in  the  text,  where  it  is  desired  more  particularly  to  trace  those 
culminating  in  existing  doctrines.  Thus  did  Reichert^  early  (1840), 
dissent  from  Schwann,  since  he  failed  to  find  the  nucleus  univer- 
sally present  in  the  yolk.  Karsten2  (1843),  published  a  dissertation 
upon  the  cell,  in  which  he  stated  that  cells  originate  without  a  pre- 
existing nucleus,  and  by  the  expansion  of  amorphous  granules  of 
organic  matter;  and  more  recently  (1863),  the  same  author  practi- 
cally reiterates  this  view,  since  he  says  that  all  "  cells  of  vegetables 
originate  as  minute  free  vesicles  in  the  fluid  contents  of  previously 
existing  cells,"  and  regards  the  nucleus  as  a  "small  tertiary  cell, 
retarded  in  its  development."^  Again,  "when  the  nucleus  is 
present,  the  origin  of  new  cells  is  quite  independent  of  it."*  In 
addition  to  the  statement  contained  in  the  text,  Henle  also  (1843), 
alleged  that  some  of  the  so-called  fibrous  tissues  were  "formed  by  the 
aggregation  of  granules  in  a  certain  way  without  the  intervention 

1  Reichert,  Das  Entwickelungslebun  im  Wirbelthlerreich.  Berlin:  1840, 
pp.  6,  9.3.         - 

2  Karsten,  De  Cella  vitale  Dissertatio.     Berlin  :  184.3. 

3  Karsten,  Ann.  and  Mag.  of  Nat.  History,  vol.  xiii,  p.  268.  London  ; 
1864. 

*  Karsten,  Ann.  and  Mag.  Nat.  History,  vol.  xiii,  p.  281. 

4* 


42  THE    CELL   DOCTRINE. 

MARTIN  BARRY,  1840. 

It  was  in  his  first  series  of  embrjological  researches, 
published  in  Part  II,  of  the  "  Philosophical  Transac- 
tions "  of  London,  for  1838,  p.  310,  that  Dr.  Martin 
Barry  declared  "that  the  germinal  vesicle  (which  he 
regarded  as  the  nucleus),  and  its  contents  constitute 
throughout  the  animal  kingdom  the  most  primitive 
portion  of  the  ovum."  In  his  second  series.  Part 
II,  1839,  in  stating  that  the  germinal  vesicle  returns 
to  the  centre  of  the  cell,  'post  coitum,  he  first  pointed 
out  that  the  nucleus  does  not  always  accompany  the 
cell  through  the  whole  vital  process  at  the  periphery 
(the  original  position  according  to  Schleiden  and 
Schwann),  but  that  it  also  passes  to  the  centre,  as 
we  now  well  know.  Here,  also,  he  declares,  but  in 
bis  third  series,  Part  II,  1840,  he  demonstrates  that 
there  arise  in  the  parent  vesicle,  two  or  more  infant 
vesicles,  the  parent  vesicle  disappearing  by  liquefac- 

of  true  nucleated  cells."!  Kolliker,2  one  of  the  foremost  exponents 
of  the  cell  doctrine  of  the  present  day,  in  1844  expressed  his  dissent 
from  the  idea  of  unity  in  the  mode  of  cell  formation,  and  states  that 
if  there  is  a  single  method  of  cell  formation  which  is  invariable, 
it  remains  to  be  discovered.  Mr.  Paget,^  so  well  known  from  his 
Lectures  on  Surgical  Pathology,  suggested  in  1846,  that  a  cell 
might  arise  in  some  other  way  than  from  a  nucleus,  since  he  had 
met  morbid  growths  composed  entirely  of  fibres,  in  which  not  a 
nucleated  cell  was  present.  Most  of  which  statements  are,  how- 
ever, reconciled  by  the  information  which  has  since  been  added  to 
our  knowledge  of  the  subject. 

1  Henle,  Traite  d'Anatomie  Qenerale.     Trad.  d'Allemand,  par  A.  J. 
Jourdain,  2  vol.     Paris  :  1843,  torn.  1,  p.  374. 

2  Kolliker,  Entwickelungsgeschichte  der  Cephalapoden.    Zurich  :   1844. 

3  Paget,  Report  on  the  Progress  of  Anatomy  and  Physiology.     Br.  and 
For.  Med.  Rev.,  July,  1846. 


THE.  CELL   DOCTRINE.  43 

tion.  And  in  his  third  series,  p.  529,  he  says,  "  The 
germinal  vesicle  does  not  burst,  or  dissolve  away,  or 
become  flattened  on  or  before  the  fecundation  of  the 
ovum  as  hitherto  supposed.  It  ceases  to  be  pellucid." 
And  on  page  531,  "  The  germinal  vesicle  tills  with 
cells,  and  these  become  filled  with  the  foundations 
of  other  cells;  so  that  the  germinal  vesicle  is  gradu- 
ally rendered  opaque." 

He  also  describes  in  this  series,  in  great  detail,  the 
mode  in  which  these  cells  are  produced  from  the 
germinal  spot,  which  he  considers  in  the  light  of  a 
nucleus  to  the  germinal  vesicle.  Part  II,  1839,  p.  360. 
And  though  the  minute  details  may  not  precisely 
accord  with  those  of  the  most  recent  observations, 
the  correct  idea  is  clearly  grasped.  In  fact,  it  may  be 
said  that  in  minuteness  of  detail  alone  does  he  differ 
from  later  observers,  and  had  he  simply  stated  that 
the  young  cells  arise  from  the  nucleus  or  nucleolus  of 
the  parent  cell,  he  would  accord  precisely  with  the 
most  recent  observers.  But  he  is,  if  possible,  even 
more  explicit  when  he  says,  "The  process  inherited 
from  the  germinal  vesicle  by  its  offspring,  reappears 
in  the  descendants  of  these.  Every  cell,  whatever  its 
minuteness,  if  its  interior  be  discerned,  is  filled  with 
the  foundations  of  new  cells,  into  which  its  nucleus 
has  been  resolved."  Again  he  says,*  "  Schleiden  has 
seen  the  nucleus  undergoing  such  changes  (division), 
but  failed  to  recognize  them."  And  finally,  in  "Phi- 
losophical Transactions"  for  1841,  pp.  207-8,  we 
have  the  following  striking  paragraphs,  which  would 
seem  also  to  correct  some  previous  errors : 

*  Barry,  Philosophical  Transac,  1840,  p.  348,  g  385. 


44  THE   CELL   DOCTRINE. 

"  §  77.  I  am  very  much  inclined  to  believe,  that  in 
the  many  instances  in  which  authors  on  ^  cells'  have 
described  and  figured  more  than  one  nucleolus  in  a 
nucleus,  there  has  been  either  an  hicipient  division 
of  the  nucleus  into  discs,  or  the  nucleus  has  consisted 
of  two  or  more  discs;  the  nucleoli  of  those  authors 
having  been  the  minute  and  highly  refracting  cavi- 
ties or  depressions  in  the  discs.  If  this  has  really 
been  the  case,  it  affords  additional  evidence,  I  think, 
that  reproduction  of  cells  hy  the  process  I  have  described — 
namely^  division  of  the  nucleus  of  the  parent  cell — is  uni- 
versal—  so  numerous  have^een  the  instances  in 
question.  I  may  refer  to  the  figures  given  by 
Schwann,  who  examined  nearly  every  tissue,  and  to 
those  of  Schleiden,  whose  observations  have  been  so 
extensive  on  plants.  I  think,  indeed,  that  many  of 
the  figures  of  Schwann  afford  evidence  of  the  divi- 
sion in  question  having  taken  place.  It  is  to  be 
recognized  in  his  delineation  of  the  cells  of  cartilage, 
cellular  tissue,  middle  coat  of  the  aorta,  muscle, 
tendon,  the  feather,  &c.  The  same  remark  is  appli- 
cable to  a  figure  given  by  Reichert  of  ciliated  epithe- 
lium cells.  Dr.  Henle  found  that  in  the  layers  of 
his  '  pflaster-epithelium '  cells,  the  nucleus,  very  dis- 
tinct in  the  lower  cells,  had  almost  disappeared  in 
those  situated  in  the  upper  part.  From  this  obser- 
vation, and  from  the  presence  of  two  nucleoli  in  some 
of  the  nuclei  figured  by  this  observer,  as  well  as  from 
the  nucleus  becoming  more  granular,  I  think  it  ex- 
tremely probable  that  these  cells  (including  those  of 
the  epidermis),  are  reproduced  by  the  process  just 
referred  to  —  division  of  the  nucleus;  additions  being 


THE   CELL   DOCTRINE.  45 

no  doubt  continuously  made  at  the  lower  part  of  the 
layer,  by  '  which  cells  previously  there  are  pushed 
farther  out.' 

"  §  83.  The  nuclei  which  various  observers  have 
found  lying  among  the  fibres  of  various  tissues,  have 
been  considered  by  them  as  the  ^remains  of  cells.' 
This  may  have  been  the  case,  but  so  far  from  think- 
ing with  those  observers,  that  the  nuclei  in  question 
were  *  destined  to  be  absorbed,'  I  am  disposed  to 
consider  that  they  were  sources  from  which  there 
would  have  arisen  new  cells." 

Without  doubt,  we  can  say,  as  did  Goodsir,*  in  the 
above  by  Martin  Barr}^  we  have  the  "  first  consis- 
tent account  of  the  development  of  cells  from  a  parent 
centre,  and  more  especially  of  the  appearance  of 
centres  within  the  original  sphere."  JS^othing  more 
definite,  or  directly  to  the  point,  could  be  desired, 
and  we  think  it  may  be  justly  said  of  Barry,  that  he 
completed  the  expression  of  the  cell  theory  inaugu- 
rated by  Schleiden  and  Schwann,  in  modifying  the 
mode  of  origin  to  conform  to  most  recent  observa- 
tion. I 

PROF.  JOHN  GOODSIR,  1845. 

In  1845,  Prof.  John  Goodsir  published  his  paper 
on  "Centres  of  I^utrition,"t  in  "Anatomical  and 
Pathological  Observations,"  in  which  he  clearly 
grasped  the  two  important  principles  of  the  modern 
Cellular  Pathology;  first,  the  activity  of  these  centres 

*  Goodsir,  Turner's  Edition  of  Anatomical  Memoirs.     Edin- 
burgh :  1868.     Note  on  p.  390. 
f  Goodsir,  op.  citat.,  p.  389. 


46  THE   CELL   DOCTRINE. 

(nuclei),  their  power  "  to  draw  from  the  capillary 
vessels,  or  from  other  sources,  the  materials  of  nutri- 
tion, and  to  distribute  them  by  development  to  each 
organ  or  texture  after  its  kind;"  second,  the  origin  of 
such  centres  or  nuclei  from  previously  existing  nu- 
clei. In  this  short  paper  of  three  pages,  are  con- 
tained, as  stated,  the  essentials  of  the  cell  doctrine 
of  Virchow,  and  as  it  has  recently  assumed  addi- 
tional interest  on  controversial*  grounds,  it  may  be 
well  to  introduce  as  much  as  bears  directly  upon  the 
subject.  "The  centre  of  nutrition  with  which  we 
are  most  familiar,  is  that  from  which  the  whole  or- 
ganism derives  its  origin — the  germinal  spot  of  the 
ovum.  From  this,  all  the  other  centres  are  derived, 
either  mediately  or  immediately;  and  in  directions, 
numbers,  and  arrangements,  which  induce  the  con- 
figuration and  structure  of  the  being.  As  the  entire 
organism  is  formed  at  first,  not  by  simultaneous 
formation  of  its  parts,  but  by  the  successive  devel- 
opment of  these  from  one  centre,  so  the  various  parts 
arise  each  from  its  own  centre,  this  being  the  orig- 
inal source  of  all  the  centres  with  which  the  part  is 
ultimately  supplied. 

"From  this  it  follows,  not  only  that  the  entire 
organism,  as  has  been  stated  by  the  authors  of  the 
cellular  theory,  consists  of  simple  or  developed  cells, 
each  having  a  peculiar  independent  vitality,  but 
that  there  is  in  addition,  a  division  of  the  ichole  into 
departments,  each  containing  a  certain  number  of 
developed   cells,  all  of  lohich  hold  certain  relations, 

*  Edinburgh  Monthly  Medical  Journal,  Feb.  and  April,  1869, 
pp.  766  and  959. 


THE    CELL    DOCTRINE.  47 

to  one  central  or  capital  cell,  around  which  they  are 
grouped.  It  would  appear  that  from  this  central  cell, 
all  the  other  cells  of  its  department  derive  their 
origin.  It  is  the  mother  of  all  those  within  its  own 
territory.  It  has  absorbed  materials  of  nourishment 
for  them  while  in  a  state  of  development,  and  has 
either  passed  them  off  after  they  have  been  fully 
formed,  or  have  arrived  at  a  stage  of  growth  when 
they  can  be  developed  by  their  own  powers. 

"Centres  of  nutrition  are  of  two  kinds, — those 
which  are  peculiar  to  the  textures,  and  those  which 
belong  to  the  organs.  The  nutritive  centres  of  the 
textures  are  in  general  permanent.  Those  of  the 
organs  are  in  most  instances  peculiar  to  their  em- 
bryonic stage,  and  either  disappear  ultimately  or 
break  up  into  the  various  centres  of  the  textures  of 
which  the  organs  are  composed. 

"  A  nutritive  centre,  anatomically  considered,  is  merely 
a  cell,  the  nucleus  of  lohich  is  the  permanent  source  of  suc- 
cessive broods  of  young  cells,  which  from  time  to  time 
fill  the  cavity  of  their  parent,  pass  off  in  certain 
directions,  and  under  various  forms,  according  to 
the  texture  or  organ  of  which  their  parent  forms  a 
part." 

Prof.  Goodsir  does  not  fail  to  state  in  the  first  para- 
graph of  his  paper,  that  with  many  of  these  centres 
anatomists  have  been  for  some  time  familiar,  but 
further  remarks  that  with  few  exceptions  they  have 
looked  upon  them  as  embryonic  structures.  He 
alludes  in  a  note  to  the  observations  of  Bowman 
and  Barry,  the  former  on  "Muscle,"  and  the  latter 
"  On  the  Corpuscles  of  the  Blood,"  in  Philosophical 


48  THE   CELL    DOCTRINE. 

Transactions,  respectively,  of  1840  and  1841,  and 
states  in  a  second  note  that  "for  the  first  consistent 
account  of  the  development  of  cells  from  a  parent 
centre,  and  more  especially  the  appearance  of  new 
centres  within  the  original  sphere,  we  are  indebted 
to  Martin  Barry."*  We  have  carefully  read  the 
references  in  each  instance.  In  Bowman's  paperf 
we  can  recognize  a  brief  reference  to  a  possible 
influence  of  the  cell  upon  nutrition,  but  none  as  to 
its  origin,  in  the  following  sentence:  "It  is,  how- 
ever, not  impossible,  that  in  all  these  cases,  there  may 
be  during  development,  and  subsequently,  a  further 
and  successive  deposit  of  corpuscles  (nuclei)  from 
which  both  growth  and  nutrition  may  take  their 
source."  That  Dr.  Barry's  paper  is  more  explicit 
has  been  shown. 

HUXLEY,  1853.| 

Allusion  has  already  been  ^made  to  Prof  Huxley 

*  Goodsir,  Anatomical  Memoirs,  vol.  ii,  p.  389,  and  note  on 
pp.  390-91. 

f  Bowman,  "Muscle,"  Philos.  Transac,  1840,  pt.  i,  p.  485. 

I  We  presume  it  will  scarcely  be  inferred  by  any  reader,  that 
the  views  of  Prof.  Huxley  here  presented  are  brought  forward  as 
those  now  entertained  by  the  learned  Professor,  and  with  which 
the  public  have  been  made  so  generally  familiar  through  his  lec- 
tureon  "  Protoplasm,"  or  the  *'  Physical  Basis  of  Life,"  delivered 
at  Edinburgh,  November  18th,  1868,  and  originally  published  in 
the  "Fortnightly  Keview  "  for  February,  1869;  but  also  largely 
republished  in  numerous  English  and  American  periodicals,  as 
well  as  in  a  separate  pamphlet,  to  be  had  of  the  publishers  of  the 
Yale  College  Courant,  New  Haven,  Conn.  To  one  closely  observ- 
ing, however,  we  think  that  these  later  views  will  appear  to  be 
foreshadowed  in  the  theory  here  given,  and  which  we  think  of 
suflBcient  historical  importance  to  justify  its  presentation  here. 


THE    CELL    DOCTRINE.  49 

in  connection  with  Wolff,  of  whose  theory  he  has 
been  the  able  exponent.  In  the  same  paper*  he 
has  given  us  his  own  views — "  conceived  in  the  spirit, 
and  not  unfrequently  borrowing  the  phraseology,  ot 
Wolff  and  Yon  Baer."  We  present  them,  as  far  as 
may  be  consistent  with  brevity,  in  his  own  w^ords: 

"Vitality,  the  faculty,  that  is,  of  exhibiting  defi- 
nite cycles  of  change  in  form  and  composition,  is  a 
property  inherent  in  certain  kinds  of  matter.  There 
is  a  condition  of  all  kinds  of  living  matter  in  which 
it  is  an  amorphous  germ — that  is,  in  which  its  exter- 
nal form  depends  merely  on  ordinary  physical  laws, 
and  in  which  it  possesses  no  internal  structure. 
Now,  according  to  the  nature  of  certain  previous 
conditions,  the  character  of  the  changes  undergone, 
or  the  different  states  exhibited — or,  in  other  words, 
the  successive  differentiations  of  the  amorphous  mass 
will  be  different. 

"  The  morphological  differentiation  may  be  of  two 
kinds.  In  the  lowest  animals  and  plants, — the  so- 
called  unicellular  organisms — it  may  be  said  to  be 
external,  the  changes  of  form  being  essentially  con- 
filled  to  the  outward  shape  of  the  germ,  and  being 
unaccompanied  by  the  development  of  any  internal 
structure. 

"But  in  all  other  animals  and  plants,  an  internal 
morphological  differentiation  precedes  or  accompa- 
nies the  external,  and  the  homogeneous  germ  becomes 
separated  into  a  certain  central  portion,  which  we 
have  called  the  endoj^last,  and  a  peripheral  portion, 

*  Huxley,  Keview  of  the  Cell  Theory.  Br.  and  For.  Med.-Chir. 
Rev.,  Oct.,  1853,  p.  305. 


50  THE    CELL    DOCTRINE. 

the  periplast.  Inasmuch  as  the  separate  existence  of 
the  former  necessarily  implies  a  cavity  in  which  it 
lies,  the  germ  in  this  state  constitutes  a  vesicle  with 
a  central  particle,  or  a  'nucleated  cell.'  There  is  no 
evidence  ivhaiever  that  the  molecular  forces  of  the  liv- 
ing matter  (the  *vis  essentialis'  of  Wolf,  or  the 
vital  forces  of  the  moderns),  are  by  this  act  of  differ- 
entiation localized  in  the  endoplast  to  the  exclusion 
of  the  periplast,  or  vice  versa.  Neither  is  there  any 
evidence  that  any  attraction  or  other  influence  is  exercised 
by  the  one  over  the  other;  the  changes  which  each  sub- 
sequently undergoes,  though  they  are  in  harmony, 
having  no  causal  connection  with  one  another,  but  each 
proceeding,  as  it  would  seem,  in  accordance  with  the 
general  determining  laws  of  the  organism.  On  the 
other  hand,  the  *vis  essentialis'  appears  to  have  es- 
sentially different  and  independent  ends  in  view,  in 
thus  separating  the  endoplast  from  the  periplast. 

"The  endoplast  grows  and  divides;  but,  except  in 
a  few^  more  or  less  doubtful  cases,  it  would  seem  to 
undergo  no  other  morphological  change.  It  fre- 
quently disappears  altogether;  but  as  a  rule  it  under- 
goes neither  chemical  nor  morphological  metamor- 
phosis. So  far  from  being  the  centre  of  activity  of 
the  vital  actions,  it  would  appear  much  rather  to  be 
the  less  important  histological  element. 

^' The  periplast,  on  the  other  hand,  which  has  hith- 
erto passed  under  the  name  of  cell  wall,  contents 
and  intercellular  substance,  is  the  subject  of  all  the 
most  important  metamorphic  processes,  whether 
morphological  or  chemical,  in  the  animal  and  plant. 
By  its  differentiation,  every  variety  of  tissue  is  pro- 


THE    CELL   DOCTRINE.  51 

duced;  and  this  difterentiation  is  the  result,  not  of 
any  metabolic  action  of  the  endoplast,  which  has  fre- 
quently disappeared  before  the  metamorphosis  begins, 
but  the  intimate  molecular  changes  in  its  substance, 
which  take  place  under  the  guidance  of  the  'vis 
essentialis,'  or,  to  use  a  strictly  positive  phrase,  occur 
in  a  definite  order,  we  know  not  why. 

"  The  metamorphoses  of  the  periplastic  substance 
are  twofold  —  chemical  and  structuraL  The  former 
(chemical),  may  be  of  the  nature  either  of  conversion, — 
change  of  cellulose  into  xylogen,  intercellular  sub- 
stance, &c.,  of  the  indifferent  tissues  of  embryos  into 
collagen,  chondrin,  &c., — or  of  deposit, — as  of  silica 
in  plants,  of  calcareous  salts  in  animals.  The  struc- 
tural metamorphoses,  again,  are  of  two  kinds,  vacuola- 
Hon  or  the  formation  of  cavities,  as  in  the  intercellu- 
lar passages  of  plants,  the  first  vascular  canals  of 
animals  —  and  fibrillaiion,  or  the  development  of  a 
tendency  to  break  up  in  certain  definite  lines  rather 
than  in  others." 

These  views  he  illustrates  by  examples  from  vege- 
table life  in  the  sphagnum  leaf,  and  from  animal  life 
in  connective  tissue  and  striped  muscle. 

As  characteristic  and  distinguishing  features  of 
this  theory,  we  desire  to  point  out,  first,  the  substitu- 
tion of  the  term  ^^ endoplast"  for  "nucleus;"  that 
of  ^'periplast''  for  "cell  wall,"  and  "intercellular," 
"substance."  Second,  the  absolutely  passive  na- 
ture of  the  "endoplast,"  which  is  neither  itself  the 
author  of  changes,  nor  the  subject  of  changes. 
Third,  the  passive  nature  as  well,  of  the  "  periplast," 
so  far  as  it  is  the  author  of  changes,  though  it  is  pre- 


52  THE    CELL   DOCTRINE. 

eminently  the  subject  of  changes,  the  seat  in  which 
changes  take  place.  And  herein,  we  believe  Huxle}^ 
to  have  been  misinterpreted  by  some  who  have 
presented  his  views  elsewhere,  as  Dr.  Beale,*  who 
represents  him  as  believing  the  periplast  active,  that 
it  is  the  efficient  agent,  that  it  sends  in  partitions, 
&c.  But  that  Prof.  Huxley  considered  it  passive  we 
believe  may  be  legitimately  inferred  from  his  text. 
As  the  seat  of  change,  however,  accomplished  not  as 
"the  result  of  any  metabolic  action  of  the  endoplast, 
but  of  intimate  molecular  changes  in  its  substance, 
which  take  place  under  the  guidance  of  the  vis  es- 
sentialis,''  the  periplast  is  differentiated  into  every 
variety  of  tissue.  Finally,  we  have  the  distinct  ad- 
mission, as  seen  in  the  sentence  last  quoted,  and  also 
throughout  the  entire  expression  of  the  theory,  of  a 
controlling,  guiding  principle,  through  which  the 
differentiation  is  accomplished.  This  principle, 
which  is  here  referred  to  as  the  "  vis  essentialis,"  is 
elsewhere  included  under  the  expressions  "  vitality," 
and  "general  determining  laws  of  the  organism." 
Though  this  admission  is  seemingly  so  at  variance 
with  the  views  of  the  same  observer  at  the  present 
date  (1870),  who  in  common  with  other  physicists 
emphatically  denies  the  existence  of  "vital  force," 
or  even  such  a  thing  as  life  itself,  yet,  as  already 
intimated,  we  deem  it  possible  to  detect  a  fore- 
shadowing of  his  more  modern  views,  in  the  follow- 


*  Beale,  Microscope  in  Medicine.  Third  Edition.  London : 
1867,  page  147.  Beale,  Structure  and  Growth  of  the  Tissues. 
London :  1865,  pp.  9,  10. 


THE   CELL   DOCTRINE.  63 

ing  paragraph  of  the  paper  whence  we  have  derived 
our  information : 

"  We  have  therefore  maintained  the  broad  doctrine 
established  by  Wolff,  that  the  vital  phenomena  are 
not  necessarily  preceded  by  organization,  nor  are  in 
any  way  the  result  or  effect  of  formed  parts,  but  that 
the  faculty  of  manifesting  them  resides  in  the  matter 
of  which  living  bodies  are  composed,  as  such — or, 
to  use  the  language  of  the  day,  that  the  vital  forces 
are  molecular  forces  J^"^ 

Huxley  moreover  says  that  the  three  botanical 
data  upon  which  Schwann's  theory  was  based,  viz. : 

1.  The  anatomical  independence  of  the  vegetable 
cell  as  a  separate  entity, 

2.  His  conception  of  the  structure  of  the  vegeta- 
ble cell,  and 

3.  Its  mode  of  development,  were  all  erroneous. 
Since  first,  he  (Huxley)  considers  that  the  fact  that 

by  certain  chemical  or  mechanical  means,  a  plant 
may  be  broken  up  into  vesicles,  corresponding  w^ith 
the  cavities  which  previously  existed  in  it,  is  of  no 
more  value  in  proving  the  independence  of  these 
vesicles,  than  the  fact  that  a  rhombohedron  of  spar, 
broken  up  with  the  hammer,  into  minute  rhombo- 
hedrons,  is  evidence  that  those  minuter  ones  were 
once  independent,  and  formed  the  larger  by  their 
coalescence. 

Second,  Schwann's  view  of  the  anatomy  of  the 
cell  was  incorrect,  since  he  regarded  the  nucleus  as 
invariably  present,  whereas  in  certain  vegetable  cells 

*  Huxley,  loc.  citat.,  p.  314. 
5* 


64  THE   CELL   DOCTRINE. 

(as  in  Hjdrodictyon,  Yaucheria,  Caulerpa,  Sphag- 
num), it  is  indubitably  absent;  and  since  he  did  not 
include  the  nitrogenous  primordial  utricle,  discov- 
ered by  Mohl,  in  1844,*  as  one  of  the  elements  of 
the  cell. 

Finally,  Schwann's  mode  of  cell-development  is 
erroneous,  having  "  been  long  since  set  aside  by  the 
common  consent  of  all  observers;"  cell-development 
always  occurring  by  division,  except  in  the  embryo 
sac  of  the  Phanerogamia,  the  sporangia  of  Lichens, 
and  of  some  Algse  and  Fungi ;  and  even  the  free 
cell-development  of  the  latter  is  quite  diiferent  from 
that  of  Schleiden  and  Schwann,  being  by  the  devel- 
opment of  a  cellulose  membrane  (periplast)  around 
a  mass  of  nitrogenous  substance  (endoplast),  which 
may  or  may  not  contain  a  nucleus. 

The  diiference  between  the  views  of  Schwann  and 
Huxley  are  best  expressed  by  the  latter  in  the  con- 
trast he  draws  between  those  of  Schwann  and  Wolf: 
"For  Schwann,  the  organism  is  a  beehive, its  action 
and  forces  resulting  from  the  separate  but  harmoni- 
ous action  of  all  its  parts.  For  Wolff'  (and  Huxley), 
it  is  a  mosaic,  every  portion  of  which  expresses  only 
the  conditions  under  which  the  formative  power 
acted,  and  the  tendencies  by  which  it  was  guided." 

The  statements  of  Prof.  Huxley  with  regard  to 
cell-development  entirely  accord  with  the  most  recent 
observations  on  the  subject,  and  are  quite  important 
to  us  in  tracing  out  the  present  state  of  the  cell  doc- 
trine. 

*  The   existence  of  the  primordial  utricle  is  denied  by  many 
botanists  of  the  present  day. 


THE   CELL   DOCTRINE.  66 


J.  hughes' BENNETT,  1866.* 

Dr.  Bennett,  of  Edinburgh,  considers  that  "the 
ultimate  parts  of  organization  are  not  cells  nor 
nuclei,  but  the  minute  molecules  from  which  these 
are  formed.  They  possess  independent  physical  and 
vital  properties,  which  enable  them  to  unite  and 
arrange  themselves  so  as  to  produce  higher  forms. 
Among  these  are  nuclei,  cells,  fibres,  and  membranes, 
all  of  which  may  be  produced  directly  from  mole- 
cules. The  development  and  growth  of  organic 
tissues  is  owing  to  the  successive  formation  of  his- 
togenetic  and  histolytic  molecules.  The  breaking 
down  of  one  substance  is  often  the  necessary  step  to 
the  formation  of  another;  so  that  the  histolytic  or 
disintegrative  molecules  of  one  period  become  the 
histogenetic  or  formative  molecules  of  another." 

Again  :  "  As  to  development,  the  molecular  is  the 
basis  of  all  the  tissues.  The  first  step  in  the  process 
of  organic  formation  is  the  production  of  an  organic 
fluid ;  the  second,  the  precipitation  in  it  of  organic 
molecules,  from  which,  according  to  the  molecular 
law  of  growth,  all  other  textures  are  derived  either 
directly  or  indirectly. "f 


*  Bennett's  Practice  of  Medicine.  Am.  Ed.  of  Wm.  Wood  & 
Co.,N.  Y.:  1866,  p.  118. 

Prof.  Bennett  has  further  elaborated  his  views  in  the  Edinburgh 
Medical  Journal,  March,  1868,  and  The  Popular  Science  Review, 
January,  1869,  but  his  conclusions  are  substantially  the  same  as 
quoted. 

t  Op.  citat.,  p.  119. 


56 


THE   CELL   DOCTRINE. 


Figs.  4,  5,  6,  7,  illustrate  these  views  amply. 


Fig.  4.      Fig.  5. 


Fig.  6. 


Fig.  7. 


Fig.  4,  Molecular  structure  of  the  scum  on  its  first  appearance,  in  a 
clear  animal  infusion.  Fig,  5,  Molecular  structure  of  the  same  six  hours 
afterwards.  The  molecules  are  separated,  and  the  long  ones  (so-called 
vibriones)  in  active  movement.  Fig.  6,  The  same  on  the  second  day. 
Fig.  7,  Filaments  (so-called  spirilla)  formed  by  aggregation  of  the  mole- 
cules, in  the  same  scum  on  the  third  and  fourth  days,  all  in  rapid  motion. 
800  diam.  linear.     (From  Bennett's  Practice.) 

Prof.  Bennett  contends,  also,  that  morbid  growths 
may  easily  be  shown  to  originate  in  a  molecular 
blastema,  though  not  to  the  exclusion  of  pre-existing 
cells.  The  accompanying  figures  are  sufficiently  ex- 
planatory. 


Fig.  8 


Fig.  10. 


Fig.  11. 


Fig.  8,  Nuclei  imbedded  in  a  molecular  blastema.  Fig.  9,  Young  fibre- 
cells  formed  by  the  aggregation  of  molecules  around  the  nuclei  of  Fig.  8. 
Fig.  10,  Cancer  cells,  one  with  a  double  nucleus.  Fig.  11,  Histolytic  or 
so-called  granule-cells,  breaking  down  from  fatty  degeneration.  250  diam. 
linear.     (From  Bennett's  Practice.) 


It  should  be  stated  also,  that  this  author  in  com- 
mon with  others  not  accepting  the  cell  doctrine  in 
its  entirety,  admits  the  production  of  cells  by  buds, 
division,  or  proliferation,  without  a  new  act  of  gene- 
ration, and  that  "  this  fact  comprehends  most  of  the 


THE   CELL   DOCTRINE.  57 

admitted  observations  having  reference  to  the  cell 
doctrine."* 

We  have  in  the  expression  of  this  theory,  a  prac- 
tical admission  of  the  spontaneous  origin  of  ani- 
mal life,  of  which  Dr.  Bennett,  in  the  paper  re- 
ferred to  in  the  Popular  Science  Review,  for  Jan., 
1869,  openly  declares  himself  the  advocate. 

Closely  allied  to  this  theory  is  the  so-called  in- 
vestment or  cluster-theory  (Umhlillungs-theorie),  de- 
scribed by  Yirchow  on  page  53,  of  Cellular  Pathology, 
(Am.  Ed.  of  Chance's  Translation);  according  to  which 
"originally  a  number  of  elementary  globules  existed 
scattered  throughout  a  fluid,  but  that  under  certain 
circumstances  they  gathered  together,  not  in  the 
form  of  vesicular  membranes,  but  so  as  to  constitute 
a  compact  heap,  a  globe  (mass,  cluster — KlUmpchen), 
and  that  this  globe  was  the  starting-point  of  all  fur- 
ther development,  a  membrane  being  formed  outside 
and  a  nucleus  inside,  by  the  differentiation  of  the 
mass,  by  apposition,  or  intussusception." 

Fig.  12. 


Fig.  12.  Diagram  of  the  Investment  (cluster)  theory.  «,  Separate 
elementary  granules,  h,  Heap  of  granules  (cluster),  c,  Granule  cell, 
with  membrane  and  nucleus. 


TODD  AND  BOWMAN,  1856. 

Notwithstanding   earlier    approximations   to   the 


*  Op.  citat.,  p.  123. 


68  THE   CELL   DOCTRINE. 

truth,  we  find  free  cell  formation  still  admitted  by  the 
eminent  authorities  Todd  and  Bowman,  as  one  mode 
of  origin  of  cells,  so  late  as  December,  1856,  though 
the  spontaneous  origin  of  organs  is  spoken  of  as  ex- 
ceedingly doubtful.  After  describing  the  elements 
of  the  ovum,  considered  in  its  entirety  as  a  nucleated 
cell,  and  referring  to  the  period  after  fecundation,  it 
is  stated,  "  At  this  period  the  embryo  consists  of  an 
aggregate  of  cells,  and  its  further  growth  takes  place 
by  the  development  of  new  ones.  This  may  be  ac- 
complished in  two  ways;  first,  by  the  development 
of  new  cells  within  the  old,  through  the  subdivision 
of  the  nucleus  into  two  or  more  segments,  and  the 
formation  of  a  cell  around  eaoh,  which  then  becomes 
the  nucleus  of  a  new  cell,  and  may  in  its  turn  be  the 
parent  of  other  nuclei ;  and,  secondly,  by  the  forma- 
tion of  a  granular  deposit  between  the  cells,  in 
which  the  development  of  the  new  cells  takes  place. 
The  granules  cohere  to  each  other  in  sejyarate  groups^  here 
and  there  J  to  form  nuclei,  and  around  each  of  these  a  deli- 
cate membrane  is  formed^  which  is  the  cell  membrane. 
The  nuclei  have  been  named  cgtoblasts,  because  they 
appear  to  form  the  cells;  and  the  granular  deposit  in 
which  these  changes  take  place  is  called  the  cytoblas- 
tema. 

"In  every  part  of  the  embryo  the  formation  of 
nuclei  and  of  cells  goes  on  in  one  or  both  of  the 
ways  above  mentioned,  and,  by  and  by,  ulterior 
changes  take  place,  for  the  production  of  the  ele- 
mentary parts  of  the  tissues."* 

*  Todd  and  Bowman,  The  Physiological  Anatomy  and  Phj'siol- 
ogy  of  Man.     Am.  Edit.,  Philadelphia:  1857,  p.  63. 


THE    CELL    DOCTRINE.  59 

Thus  did  physiologists  adhere  to  the  original  free 
cell  formation  of  Schleiden  and  Schwann.  Singu- 
larly, Dr.  Carpenter,*  who  expressly  states,  in  his 
Manual  of  Physiology,  edition  of  1865,  that  he  has 
been  led  to  the  view  of  Professor  Beale  by  com- 
parison of  the  results  of  the  recent  inquiries  of  sev- 
eral British  and  Continental  histologists  with  those 
of  his  own  studies,  says,  a  few  pages  further  on  (p. 
150),  "]^ew  cells  may  originate  in  one  of  two  prin- 
cipal modes;  either  directly  from  a  previously  exist- 
ing cell,  or  by  an  entirely  new  process  in  the  midst 
of  an  organizable  blastema."  He  then  proceeds  to 
give  the  two  methods  in  detail,  without  in  any  way 
denying  the  latter. 

viRCHOW,  1858. 

Less  than  two  years  later,  August  20th,  1858,  Prof. 
Yirchow  published  his  '*  Cellular  Pathology  as  based 
upon  Physiological  and  Pathological  Histology." 
According  to  him,  the  cell  is  the  only  possible  start- 
ing-point for  all  biological  doctrines.  This  cell  can 
only  originate  from  a  previously  existing  cell,  taking 
its  primary  origin  from  the  ovum,  and  the  Harveian 
maxim  omne  vivum  ex  ovo,  becomes  in  its  special 
application,  omnis  cellula  e  celluld.  This  is  true  of  all 
physiological  and  pathological  processes  in  the  vege- 
table and  animal.  In  all  editions  of  "  Cellular 
Pathology"  which  we  have  met,  the  typical  cell  is 
described   as  consisting  essentially  of  "cell  wall," 

*  Carpenter,  Manual  of  Physiology,  London :  1865.  Note  on 
p.  14. 


60  THE    CELL    DOCTRINE. 

"cell  contents,"  and  "nucleusV  the  "nucleolus," 
though  usually  met  in  fully  developed  older  forms, 
is  not  considered  an  essential  constituent  of  the 
cell.  The  object  of  the  "  nucleus,"  according  to  * 
Yirchow,  is  entirely  connected  with  the  life  of  the 
cell,  that  which  maintains  it  as  an  element  and 
which  insures  its  reproduction.  While  to  the  "cell 
co>utents"  over  and  above  the  nucleus,  that  is  the 
"  residual  cell  contents,"  is  due  the  function  of  the 
cell,  that  to  which  is  due  the  contractility  of  mus- 
cle, the  neurility  and  sensation  of  nerve,  and  the 
secretory  office  of  the  gland  cell.* 

To  secure  the  universal  application  of  the  cell 
doctrine,  it  becomes  necessary  to  eliminate  from  the 
vegetable  cell,  the  external  non-nitrogenous  mem- 
brane known  as  cellulose,  and  restrict  it  to  the  nitro- 
genized  portion  comprised  in  the  primordial  utricle  as 
the  proper  cell  wall,  and  in  the  protoplas7nic  contents 
of  the  cavity  as  the  proper  cell  contents,  which  con- 
tain also  the  nucleus.  "  It  is  only  when  we  adhere 
to  this  view  of  the  matter,  when  we  separate  from 
the  cell  all  that  has  been  added  to  it  as  an  after-de- 
velopment, that  we  obtain  a  simple,  homogeneous, 
extremely  monotonous  structure,  recurring  with  ex- 
traordinary frequency  in  living  organisms."! 

More  recently,  however,  Virchow  is  reported  as 
not  regarding  the  "cell  wall"  as  an  essential  part  of 
the  cell,  as  stated  in  Cellular  Pathology;  but  that  a 

*  Virchow,  Cellular  Pathology  as  based  upon  Physiological  and 
Pathological  Histology.  Second  Edition.  Translated  by  Frank 
Chance,  MB.,  «&c.     Am.  Edition,  Philadelphia:  1863,  p.  37. 

t  Op.  cit.,  pp.  31,  34. 


THE   CELL   DOCTRINE.  61 

nucleus  surrounded  hy  a  molecular  blastema  was  sufficient 
to  constitute  a  cell;  then  he  says  that  the  outer  part  of 
this  cell  blastema  consolidates  and  forms  a  cell  wall 
as  Beale  has  shown,  and  that  this  takes  place  in  the 
amoeba  when  placed  in  water.* 

As  thus  defined,  the  cell  is  the  seat  of  pathologi- 
cal and  physiological  processes  rather  than  the  blood, 
or  the  nerves.  The  cell  is  active, — the  ultimate  mor- 
phological element  in  which  there  is  any  manifesta- 
tion of  life,  and  beyond  which  the  seat  of  real  action 
cannot  be  removed.  Hence  the  term  Cellular  Pathol- 
ogy rather  than  humoral,  or  neural,  or  solidistic.  The 
so-called  exudations  are  not  such  in  the  strict  sense  of 
the  term,  and  the  cells  which  they  contain,  whether 
of  pus  or  organizable  lymph,  are  the  result  of  pro- 
liferation of  previously  existing  cells.  Even  "fibrin, 
wherever  it  occurs  in  the  body  external  to  the  blood, 
is  not  to  be  regarded  as  an  excretion  from  the  blood, 
but  as  a  local  production,"  resulting  from  the  activity 
of  the  cells  of  the  tissue  in  which  it  is  found,  and  con- 
veyed to  the  surface  by  the  transudation  oithQ  serous 
fiuids  alone. t  In  the  above  statements  we  have  the 
first  distinctive  feature  of  Yirchow's  theory. 

Again,  since  every  organized  body  is  usually  made 
up  of  a  number  of  these  cells,  each  independent  in 
itself,  yet  combined  and  arranged  for  the  attainment 
of  a  special  end,  and  therefore  mutually  dependent, 
there  result  certain  communities  or  cell  territories 
into  w^hich  the  body  is  portioned  out  by  Yirchow. 

*  Letter  from  Berlin,  in  Edinburgh  Medical  Journal,  February, 
1865. 
f  Virchow,  op.  cit.,  pp.  435-6. 

6 


62  THE    CELL    DOCTRINE. 

But  not  only  is  the  relation  of  these  cells  to  each 
other  and  to  the  central  cell  whence  they  took  their 
origin  mutually  dependent,  but  in  many  animal  tis- 
sues, at  least,  we  have  the  so-called  intercellidar  sub- 
stance in  a  certain  definite  manner  dependent  upon 
the  cell  or  cells  which  it  surrounds,  "so  that  certain 
districts  belong  to  one  cell  and  certain  others  to 
another."  Especially  is  this  the  case  in  pathological 
processes,  where  sharp  boundaries  may  often  be  drawn 
between  cell  territories.  Herein  have  we  the  second 
distinguishing  character  of  Virchow's  theory. 

There  are  also  a  third  and  fourth  distinctive  fea- 
tures. It  has  already  been  explained  that  the  prin- 
ciple of  the  theory  of  Schleiden  and  Schwann  lay  in 
this,  that  every  tissue,  healthy  or  morbid,  results 
from  the  apposition  of  cells,  and  that  this  principle 
is  still  observed  as  correct,  the  mode  of  origin  of  the 
primary  cell  being  alone  the  object  of  dispute.  Ac- 
cording to  Virchow,  however,  it  is  a  special  cell 
which  becomes  the  starting-point  of  physiological 
and  pathological  processes,  and  by  its  various  meta- 
morphoses constitutes  the  healthy  or  morbid  tissue, 
excepting  epithelial  formations.  This  cell  is  the 
so-called  connective  tissue  corpuscle,  or  cell  of  the  con- 
nective tissue,  which,  according  to  Virchow,  is  a  cell 
with  all  its  essential  constituents  (cell  wall,  cell  con- 
tents, and  nucleus),  and  not  a  nucleus  alone,  as  orig- 
inally described  by  Schwann,  and  later  by  Henle* 
and    Landois.f     From    the   well-known   universal 

*  Henle,  Bericht  uber  die  Fortschritte  d.  Physiol.,  1859;  1866, 
p.  41. 
f  Landois,  Zeits.  f.  wiss.  Zool.,  Bd.  xvi,  p.  1. 


THE    CELL   DOCTRINE. 


63 


prevalence  of  connective  tissue,  this  view  receives 
support.  Thus,  it  is  from  the  connective  tissue 
corpuscles  of  the  soft,  silk-like  connective  tissue,  so 
universally  present  in  muscle,  that  the  muscular  fas- 
ciculi are  primarily  developed.  It  is  from  these  that 
nerve  fibres  take  their  origin.  It  is  by  the  rapid 
proliferation  of  these  corpuscles  that  pus  is  formed, 

Fig.  13. 


Fig.  13.  Purulent  granulation  from  the  subcutaneous  tissue  of  a  rab- 
bit, round  about  a  ligature,  a.  Connective  tissue  corpuscles,  b,  Enlarge- 
ment of  the  corpuscles  with  division  of  the  nuclei,  c,  Division  of  the  cells 
(granulations),     d,  Development  of  the  pus  corpuscles.       x300.     (From 

Virchow.) 

Fig.  14. 


Fig.  14.  Interstitial  purulent  inflammation  of  muscle  in  a  puerperal 
woman,  m  m,  Primitive  muscular  fibres,  i  i,  Development  of  pus  cor- 
puscles by  means  of  the  proliferation  of  the  corpuscles  of  the  interstitial 
connective  tissue.     x280.     (From  Virchow.) 


64  THE   CELL   DOCTRINE. 

(Figs.  13  and  14);  it  is  from  the  perverted  growth 
and  development  of  these  that  tubercle  and  cancer 

Fig.  15. 


Fig.  15.  Development  of  cancer  from  connective  tissue  in  carcinoma 
of  the  breast,  a,  Connective  tissue  corpuscles,  b,  Division  of  the  nuclei. 
c,  Division  of  the  cells,  d,  Accumulation  of  the  cel,ls  in  rows,  e,  Enlarge- 
ment of  the  young  cells  and  formation  of  the  groups  of  cells  (foci,  Zellen- 
heerde),  which  fill  the  alveoli  of  cancer.  /,  Further  enlargement  of  cells 
and  groups,  g,  The  same  development  seen  in  transverse  section.  (From 
Virchow.) 

arise  (Fig.  15),  and  similarly  all  pathological  new 
formations.  None  of  these  products  are  exudations 
from  the  blood,  according  to  Virchow.  They  are 
entirely  local  in  their  origin.  In  these  views  he  is 
supported  by  the  majority  of  German  observers. 

Another  mode  of  formation  of  pus  is  however  ad- 
mitted by  Yirchow,  in  the  growth  and  development 
of  new  cells  in  epithelium,  whether  in  cuticle  or  mu- 
cous membranes.  Whether  forms  of  suppuration 
exist  which  may  be  referred  to  muscular,  nervous, 
and  capillary  elements,  he  considers  doubtful. 

A  fourth  and  final  distinctive  feature  of  Virchow's 
views,  concerning  which  there  is  less  unanimity,  even 
among  German  histologists,  is  his  peculiar  system  of 


THE    CELL   DOCTRINE. 


65 


canals  or  tubes,  produced  by  the  anastomosis  of  one 
cell  with  another,  and  which  he  considers  must  be 
classed  with  the  great  canalicular  system  of  the  body, 
as  forming  a  supplement  to  the  blood  and  lymphatic 
vessels,  and  as  filling  up  the  vacancy  left  by  the  old 
vasa  serosa  which  do  not  exist.*    (See  Fig.  16.)    Of 

Fio.  16. 


^^^ 


Fig.  16.  Connective  tissue  from  the  embryo  of  a  pig  after  long-contin- 
ued boiling.  Large  spindle-shaped  cells,  connective  tissue  corpuscles 
(Bindegewebesktirperchen),  some  isolated  and  some  still  imbedded  in  their 
basis  substance,  and  anastomosing  one  with  the  other.  Large  nuclei  with 
their  membrane  detached  ;  cell  contents  in  some  cases  shrunken.  x350. 
(From  Virchow.) 

this  system  he  also  considers  the  cord-like  fibres  of 
yellow  elastic  tissue  as  forming  a  part.f  These  he 
considers,  with  Donders,t  as  originating  by  a  trans- 


*  Virchow,  op.  citat.,  p.  76. 
f  Virchow,  op.  citat.,  p.  133,  a.  f. 
X  Bonders,  Siebold  und  Kdlliker's  Zeitschrift,  Bd.  iii. 
6* 


66  THE   CELL   DOCTRINE. 

formation  of  the  connective  tissue  corpuscles  them- 
selves. He  says,  "  The  transformation  of  these  latter 
into  the  former,  can  gradually  be  traced  with  such 
distinctness,  that  there  remains  no  doubt,  that  even 
the  coarser  elastic  fibres  directly  result  from  a  chem- 
ical change  and  condensation  of  the  walls  them- 
selves.* Where  originally  there  lay  a  cell,  provided 
with  a  delicate  membrane  and  elongated  processes, 
there  we  see  the  membrane  gradually  increasing  in 
thickness  and  refracting  the  light  more  strongly, 
whilst  the  proper  cell  contents  continually  decrease 
and  finally  disappear. 

Fig.  17. 


Fig.  17.  Elastic  networks  and  fibres  from  the  subcutaneous  tissue  of 
the  abdomen  of  a  woman,  a  a,  Large  elastic  bodies  (cell  bodies),  with 
numerous  anastomosing  processes,  b  b,  Dense  elastic  bands  of  fibres  on 
the  border  of  larger  meshes,  c  c,  Moderately  thick  fibres  spirally  coiled 
up  at  the  end.  dd,  Finer  elastic  fibres,  at  e  with  more  minute  spiral  coils. 
x300.     (From  Virchow.) 

"  The  whole  structure  becomes  in  this  way  more 


*  Virchow,  op.  citat.,  p.  133, 


THE    CELL   DOCTRINE.  67 

homogeneous,  and  to  a  certain  extent  sclerotic,  and 
acquires  an  incredible  power  of  resisting  the  influ- 
ence of  reagents,  so  that  it  is  only  after  long-contin- 
ued action  that  even  the  strongest  caustic  substances 
are  able  to  destroy  it,  whilst  it  completely  resists  the 
caustic  alkalies  and  acids  in  the  degree  of  concentra- 
tion usually  employed  in  microscopical  investigation. 
The  farther  this  change  advances,  the  more  does  the 
elasticity  of  the  parts  increase,  and  in  sections  we 
usually  find  these  fibres,  not  straight  or  elongated, 
but  tortuous,  curled  up,  spirally  coiled,  or  forming 
little  zigzags  (Fig.  17,  c,  e).  These  are  the  elements 
which  by  virtue  of  their  great  elasticity,  cause  retrac- 
tion in  those  parts  in  which  they  are  found  in  con- 
siderable quantity,  as,  for  example,  in  the  arteries. 
The  fine  elastic  fibres,  which  are  those  which  possess 
the  greatest  extensibility,  are  usually  distinguished 
from  the  broader  ones,  which  certainly  do  not  pre- 
sent themselves  in  tortuous  forms.  As  regards  their 
origin,  however,  there  seems  to  be  no  diflference  be- 
tween the  two  kinds;  both  are  derived  from  the 
connective  tissue  cells,  and  their  subsequent  arrange- 
ment is  only  a  reproduction  of  the  original  plan. 
In  the  place  of  a  tissue,  consisting  of  a  basis  sub- 
stance and  anastomosing  reticulated  cells,  there 
afterward  arises  a  tissue  with  its  basis  substance 
mapped  out  by  long  elastic  networks  with  extremely 
compact  and  tough  fibres."  This  may  be  looked 
upon  as  the  least  well-determined  of  the  important 
points  of  Virchow's  doctrine,  though  most  German 
histologists  also  favor  it.     Among  these  may  be 


68  THE   CELL   DOCTRINE. 

classed  Kolliker,*  C.  Q.  Weber,t  Leydig,t  Fried- 
reich,§  His,||  Donders,!  Wittich,**  Bottcher,tt  Bill- 
roth,! J  and  Strieker.  They  are  opposed  by  Schwann, 
Reichert,  and  Henle,  and  find  little  favor  among 
English  and  American  histologists. 

A  part  of  this  system,  also,  according  to  Virchow, 
are  the  so-called  dentinal  tubules,  the  lacunae  and 
canaliculi  of  bone,  even  the  continuity  traced  by 
Gerlach,§§  between  the  ciliated  cells  of  the  aqueduct 
of  Fallopius;  that  by  Heidenhain||||  and  BriickeTf^f 
between  the  lacteals  and  cylinder  epithelium  of  the 
intestinal  villi  of  the  rabbit,  by  means  of  corpuscles 
of  connective  tissue;  in  the  epithelium  of  the  endo- 
cardium by  Luschka;***  and  the  results  of  similar 
observations  by  Eckhart,ttt  Billroth, J JJ  and  Fried- 
reich. §§§ 

*  Kolliker,   Manual  of  Human  Microscopic  Anatomy,  p.  41. 
1860.     Also  recent  paper  in  which  he  completely  assents  to  Vir- 
chow's  views,  according  to  N.  Y.  Q.  J.  Pschy.  Med.,  July  1869. 
f  Weber,  C.  O.,  Virchow's  Archiv,  Bnd.  xiii-xv. 
J  Leydig,  Handbuch  der  Histologic.  1856. 
§  Friedreich,  Virchow's  Archiv,  Bd.  xv. 
II  His,  Beitrage  zur  Normalen  in  Pathol.  Histol.  d. Cornea. 
^  Bonders,  loc.  citat. 
**  Wittich,  Virchow's  Archiv,  Bd.  ix. 
ff  Bottcher,  Virchow's  Archiv,  Bd.  xiii. 

J  J  Billroth,  in  Beitrage  zur  Pathol.  Histol.,  1858,  admits  all  but 
the  tubular  nature  of  the  processes. 
g§  Gerlach,  Mikrosk.  Studien.  1858. 

nil  Heidenhain,Moleschott's  Untersuchungen,Bd.  iv,  1858,p.  251. 
^^  Briicke,  Moleschott's  Untersuchungen,  Bd.  viii,  1862,  p.  495. 
***  Luschka,  Virchow's  Archiv,  Bd.  ix,  p.  569. 
Iff  Eckhart,  Beitrage  Anat.  und  Physiol.  1855. 
Ill  Billroth,  Mailer's  Archiv,  1858. 
§g§  Friedreich,  loc.  citat.,  p.  538. 


THE   CELL   DOCTRINE.  69 

The  other  fibrous  element  of  areolar  or  connec- 
tive tissue,  which  forms  the  mass  of  its  bulk,  the 
pure  white  fibrous  or  waving,  does  not,  according  to 
Virchow,  have  its  origin  in  cells,  but  is  a  modifica- 
tion of  a  previously  homogeneous  intercellular  sub- 
stance, deposited  between  the  cells, — a  view  which 
in  its  glaring  departure  from  the  primary  proposition 
that  the  cell  is  the  starting-point,  and  that  every 
tissue  is  composed  of  cells  or  some  modification  ot 
cell  forms,  presents  one  of  the  few  inconsistencies 
traceable  in  the  theory  of  Virchow. 

We  think  it  proper,  in  a  historical  memoir  of  this 
kind,  to  refer  to  some  severe  critical  remarks  which 
appeared  in  the  Edinburgh  Medical  Journal  of  Feb- 
ruarj^  and  April,  1869,  in  which  Prof.  Yirchow  is  ac- 
cused of  appropriating  the  observations  of  Prof.  Good- 
sir  as  his  own.  That  there  are  points  in  common, 
it  will  be  recollected,  and,  also,  that  these  are  first, 
the  invariable  origin  of  cells  from  previously  existing 
cells,  and  second,  the  division  of  the  tissues  into  cell 
territories.  E'ow  on  the  one  hand  we  deem  that  the 
dedication  of  Yirchow's  volume  to  Prof.  Goodsir  is 
as  handsome  an  accredit  as  could  possibly  be  given 
for  whatever  of  common  there  may  be  in  the  writings 
of  the  two  authors,  and  on  the  other  hand  we  have 
seen  that  Martin  Barry  is  acknowledged  even  by 
Goodsir,  to  be  the  author  of  the  "  first  consistent 
account  of  the  development  of  cells  from  a  parent 
centre."  The  idea  of  cell  territories  seems,  however, 
to  have  originated  with  Goodsir,  nor  do  we  believe, 
for  the  reason  stated,  that  Virchow  intended  to  usurp 
his  prerogative.     The  merit  of  Virchow  consists  in 


70  THE   CELL    DOCTRINE. 

his  application  by  actual  demonstration  of  the  first 
of  these  points  to  so  large  a  variety  of  physiological 
and  pathological  processes,  to  which  is  added  original 
conception  in  the  prominence  given  to  the  connec- 
tive tissue  corpuscle  and  the  canalicular  system, 
whatever  may  be  the  truth  with  regard  to  either. 


SARCODE  OF  DUJARDIN.     PROTOPLASM  OF  MAX  SCHULTZE. 

1835-61. 

Dujardin*  had,  in  1835,  discovered  in  the  lower  ani- 
mals a  living,  moving,  contractile  substance,  which 
he  called  sarcode.  The  peculiar  appearances  of  this 
substance  attracted  the  attention  of  many  observers, 
among  whom  were  Meyen,f  Huxley,  Max  Schultze, 
John  Muller,  and  others,  who  thought  it  peculiar  to 
the  lower  animals,  and  there  was  assigned  to  it  a 
property  of  "  irritability  without  nerves."| 

The  observation  of  Siebold,§  that  the  yolk  glob- 
ules of  Planaria  exhibit  contractions  and  expansions, 
which  with  suitable  care  continue  for  hours,  and  the 
discoveries  which  followed  of  similar  movements 
and  changes  in  form,  led  Kolliker||  to  express  the 
conjecture  that  the  contents  of  all  cells  are  contractile. 
Virchowf  attributed  the  movements  to  a  contractile 

*  Dujardin,  Ann.  d.  Sciences  Nat.,  torn,  iii  et  v. 
f  Meyen,  Einschlagige  Liter,  in  E.  Iljeckel's  Die  Kadiolaren. 
1862. 

J  Schultze,  Max,  Organis.  d.  Polythalamien.    1854. 
§  Siebold,  Froriep's  Notizen,  Nr.  380,  p.  85. 
II  Kolliker,  Wurzb.  Verh.,  Bd.  viii. 
5[  Virchow,  Archiv,  Band  v. 


THE   CELL    DOCTRINE.  71 

substance.  Leyclig*  considered  the  movements  of 
the  yolk  globules  as  phenomena  of  life,  and  Kuhnef 
had  studied  physiologically  and  chemically,  sarcode 
and  muscular  tissue,  and  compared  the  irritability 
and  changes  after  death,  of  both.  But  all  consid- 
ered sarcode  as  something  different  from  the  ani- 
mal cell,  as  a  body  sui  generis.  Max  Schultze,  in 
1861,1  had  first  shown  this  analogy  between  sarcode 
and  the  contents  of  the  animal  cell,  and  that  the  en- 
tire infusorial  world,  simple  or  compound,  is  made 
up  of  cells,  thus  extending  the  typical  formative 
element  of  Schwann  to  the  entire  organized  creation. 
So  much  for  the  relation  of  sarcode  to  the  animal 
cell. 

The  comparison  between  sarcode  and  the  proto- 
plasm of  plants  was  undertaken  by  Unger,§  E. 
Brucke,||  E.  Hseckel.t  Max  Schultze,**  and  W. 
Kuhne,tt  and  by  their  efforts,  according  to  Strick- 
er,||  our  knowledge  of  the  independent  life  of  the 
cell  was  extended,  in  a  very  short  space  of  time,  fur- 
ther than  in  the  twenty  years  previous. 

Unger§§  (1855),  had  been  struck  with  the  close 

*  Leydig,  Handbuch  der  Histologie.  1856. 
f  Kuhne,  Mull.  Archiv.,  1859,  p.  817. 
X  Schultze,  Max,  Mull.  Archiv,  1861,  p.  17. 
^  Unger,  Anatomie  und  Ph3^siology  d.  Pflanzen.     1855. 
II  Briicke,  E.,  Elementarorganism,  Wien  Sitzungsb.     1861. 
TJ  Hteckle,  E.,  Radiolaren.     1862. 
**  Schultze,  Max,  Protoplasm  der  Rhizopoden  und  der  Pflanz- 
enzellen.    1863. 
fl  Kuhne,  W.,  Protoplasm  und  die  Contractilitat.    Lpzg.:  1864. 
XI  Strieker,   S.,  Handbuch   der   Lehre  von  den   Geweben  des 
Menschen  undder  Thiere.    Leipzig:    1868,  p.  3. 
gg   Unger,  op.  citat,  p.  280. 


72  THE    CELL    DOCTRINE. 

similarity  of  the  mobile  phenomena  of  the  Poljtha- 
lamise  with  those  of  the  processes  of  protoplasm 
stretched  across  the  cavity  of  many  vegetable  cells. 
Although  he  had  not  personally  investigated  the 
former,  he  became  convinced  from  Schultze's  de- 
scription, that  a  resemblance  amounting  to  identity 
existed  between  their  movements  and  the  protoplasm 
streams  of  vegetable  cells."* 

Shortly  before  this,  Pringsheimf  (1854),  showed 
that  no  such  membrane  as  a  primordial  utricle  ex- 
isted, but  that  all  within  the  cellulose  wall  of  the  liv- 
ing vegetable  cell  was  protoplasm  and  cell  fluid,  how- 
ever complex  its  composition. 

"He  admitted  that  in  the  cortical  layer  of  the  pro- 
toplasma  a  distinct  arrangement  into  layers  often 
occurred,  and  these  he  distinguished  as  the  cutane- 
ous and  granular  layers  of  the  protoplasma,  but  he 
denied  that  the  primordial  utricle  could  be  difteren- 
tiated  as  a  membrane  from  the  subjacent  protoplasm. 
If,  in  animal  cells,  partly  from  their  relatively  small 
size,  and  partly  from  their  greater  average  wealth  in 
protoplasma,  it  is  more  rarely  possible  to  make  a 
sharp  demarcation  between  a  cortical  layer  of  pro- 
toplasm and  a  cell  fluid,  there  nevertheless  exists  a 
difference  in  the  constitution  of  the  former,  such 
that  a  cutaneous  layer,  destitute  of,  or  scantily  sup- 
plied with  granules,  incloses  the  remaining  more 
granular  material.     The  white  blood-cell  may  serve 

*  DuflSn,  A.  B.,  On  Protoplasm.  Quart.  Jour.  Mic.  Sci.,  N.  S., 
vol.  iii,  1863,  p.  252. 

■)-  Pringsheim,  Untersuchungen  iiber  d.  Bau.  u.  d.  Bildung  d. 
Pflanzenzelle.     1854. 


THE   CELL   DOCTRINE.  73 

as  an  example.  This  is,  however,  very  different  from 
a  proper  membrane."* 

The  name  protoplasm  for  the  contents  of  the  ani- 
mal cell  had  already  been  brought  into  use  by 
Kemak. 

Leydig,t  iu  1856,  claimed  for  the  contents  of  the 
cell  a  higher  dignity  than  for  the  membrane  or 
cell  wall.  He  claimed  that  a  cell  was  but  protoplasm 
(klumpchen-substanz)  inclosing  a  nucleus.  The  cell 
membrane,  according  to  him,  was  simply  the  hard- 
ened periphery  of  the  substance  of  the  cell. 

To  Max  Schultze,  however,  belongs  the  credit  of 
having  fully  overturned  the  vesicular  idea  of  cells. 
In  1861,t  he  insisted  upon  some  modification  of  pre- 
vailing views,  respecting  the  relation  of  cell  wall  to 
cell  contents,  and  contended  for  a  higher  position 
for  that  part  of  the  cell  corresponding  to  the  pro- 
toplasm of  Von  Mohl  (that  within  the  so-called 
primordial  utricle),  and  showed  how  a  careful  study 
of  the  phenomena,  presented  by  the  pseudopodia 
extended  b}^  the  various  Rhizopods,  might  aid  in 
clearing  up  the  life  of  the  elements  of  the  cell. 

He  also  defined  the  cell  as  ''protoplasm  surround- 
ing a  nucleus.' '  The  importance  of  this  definition, 
as  stated  by  Stricker,§  lay  not  so  much  in  the  fact 
that  many  cells  were  denied  a  cell  wall,  as  that  the  so- 
called  cell  contents  could  now  be  made  to  harmonize 

*  Duffin,  loc.  citat.,  p.  252. 
f  Leydig,  op.  citat. 

X  Schultze,  Max,  Ueber  Muskelkorperchen,  in  Reichert  and  Du- 
bois Reymond's  Archiv,  1861. 
§  Strieker,  op.  citat.,  p.  5. 

7 


74  THE   CELL   DOCTRINE. 

with  the  animal  primordial  substance  or  sarcode. 
Schultze  illustrates  his  definition  by  the  embryo  cells 
resulting  from  the  segmentation  of  the  ovum,  as 
typical  cells,  which  are  thus  composed  of  protoplasm 
surrounding  a  nucleus,  which  nucleus,  as  well  as  pro- 
toplasm, are  products  of  like  constituent  parts  of 
another  similar  cell.  "  The  cell  leads  in  itself  an 
independent  life  of  which  the  protoplasm  is  espe- 
cially the  seat,  although  to  the  nucleus  also  undoubt- 
edly falls  a  most  important,  though  not  yet  precisely 
determined  role.  Protoplasm  is  for  the  most  part  no 
further  distinct  than  that  it  will  not  commingle  with 
the  surrounding  medium,  and  in  the  peculiarity  that 
with  the  nucleus  it  forms  a  unit.  Upon  the  surface 
of  the  protoplasm,  there  may  form  a  membrane, 
which,  although  derived  from  it,  may  be  chemically  differ- 
ent, and  the  assertion  that  it  is  the  beginning  of  a  retro- 
gression may  be  defended,  A  cell  with  a  membrane  can 
not  divide  itself,  unless  the  protoplasm  within  the 
membrane  divides  itself.  A  cell  within  a  membrane 
chemically  different  from  protoplasm,  is  like  an  en- 
cysted infusorium."* 

Bruckef  went  even  further  in  his  definition,  and 
said  that  it  was  not  shown  that  the  nucleus  even  is 
an  essential  element  of  the  cell.  In  proof  of  which 
he  adduces  the  cells  of  cryptogams.  Facts  in  justi- 
fication of  Brucke's  doubt  are  adduced  by  StrickerJ 
in  the  discovery  by  Max  Schultze,§  in  the  Adriatic 

*  Schultze,  Max,  Protopl.  d.  Khizopoden.    Leipzig:  1863. 
•}-  Briicke,  E.,  Die  Elementarorganismen,  p.  18-22,    1861. 
%  Strieker,  op.  citat.,  p.  6. 
§  Schultze,  Max,  Organis.  d.  Polythalam.     1854. 


'       THE   CELL   DOCTRINE.  75 

Sea,  of  a  non-nucleated  amoeba,  by  Hseckel*  in  the 
Mediterranean,  of  a  non-nucleated  protozoon  (Pro- 
togenes  primordialis),  and'by  Cienkowskyf  of  two 
non-nucleated  monads,  namely,  Monas  amyli  and 
Protomonas  amyli.  Hseckel  says  of  his  Protogenes 
primordialis  that  it  multiplies  by  division.  Strieker's  J 
own  observations  on  the  fecundated  egg  of  the  frog, 
incline  him  to  adopt  the  view  of  Briicke,  and  to  omit 
the  nucleus  in  a  theory  of  elementary  organization. § 
With  these  general  considerations  in  the  history 
of  "  protoplasm,"  we  are  the  better  prepared  to  take 
up  the  theory  of 

DR.  BEALE,  1861. 

In  April  and  May,  1861,  Prof.  Lionel  S.  Beale 
delivered  the  lectures  before  the  Royal  College  of 
Physicians,  of  London,  in  which  he  promulgated 

*  Hseckel,  Zeitschr.  f.  w.  Zoolog.,  1865,  Bd.  xv. 

f  Cienkowsky,  Max  Schultze's  Archiv,  1865. 

J  Strieker,  op.  citat. 

§  In  a  recent  paper  by  Prof.  Hseckel,!  he  states  that  the  "pro- 
toplasm theory"  was  brought  forward  in  its  elementary  form  by 
Cohn,2  in  1850,  and  by  Unger,  in  1853,  It  was  further  developed 
in  1858,  and  finally  completely  established  in  1860,  by  Max  Schultze. 
Hseckel  also  considers  that  by  no  phenomena  is  the  correctness  of 
this  theory  so  thoroughly  proved,  and  at  the  same  time  in  so  sim- 
ple and  unassailable  a  manner,  as  by  the  vital  phenomena  of  the 
Monera,  by  the  processes  of  their  nourishment  and  reproduction, 
sensitiveness  and  motion,  which  entirely  proceed  from  one  and  the 
same  very  simple  substance,  a  true  "primitive  slime." 

1  Ilaeckel,  Ernst,  Monograph  on  the  Monera,  and  Remarks  on  the  Pro- 
toplasm Theory.     Q.  Jour.  Mic.  Sci.,  Apr.,  July,  and  Oct.,  1869. 

2  Cohn,  F.,  Nachtrage  zur  Naturgeschichte  des  Protococcus  pluvialis ; 
Nova  Acta  Ac.  Leop.  Carol.,  vol.  xxii,  pars  ii,  p.  605.     1850. 


76  THE   CELL   DOCTRINE. 

the  views  which  have  since  been  further  elaborated 
and  become  permanently  associated  with  his  name. 
These  views  were  published,  in  part,  in  Beale's 
"Archives  of  Medicine,"  and  in  September,  1861, 
in  a  volume  "  On  the  Structure  of  the  Simple  Tis- 
sues of  the  Human  Body,"  in  the  preface  to  which 
he  says  "  he  thinks  it  right  to  state  that  the  conclu- 
sions which  have  now  assumed  a  definite  form  have 
gradually  grown  upon  him  during  the  course  of  ob- 
servations, extending  over  a  period  of  several  years. 
In  fact  some  of  the  drawings  in  this  volume,  and 
others  wdiich  have  been  published  elsewhere,  equally 
favorable  to  this  view,  were  made  long  before  any 
specific  theory  had  been  arrived  at." 

The  "ce?^,"  or  ^'elementary  fart^^  as  Dr.  Beale  pre- 
fers to  call  it,  is  composed  of  matter  in  two  states, 
matter  which  \%  forming^  and  matter  which  is  formed; 
matter  which  has  the  power  of  groioing  by  producing 
matter  like  itself  out  of  pabuhmi  or  food,  and  matter 
which  possesses  no  such  power,  but  results  from  the 
death  of  the  forming  matter.  The  former  is  known 
as  germinal  or  living  matter,  the  latter  as  formed  mat- 
ter. The  former,  in  varying  quantity  in  diflerent 
cells,  is  central  in  its  situation  (see  Plate,  Fig.  17), 
and  includes  what  has  been  called  by  others  nucleus, 
cell  contents,  protoplasm,  endoplast.  The  latter,  also 
present  in  difterent  quantity  in  diflferent  cells,  is  pe- 
ripheral (Plate,  Fig.  17),  and  includes  what  is  known 
as  cell  wall,  periplast,  intepcelldlar  substance,  and 
products  of  secretion. 

In  its  structural  characters,  germinal  matter  is  soft, 
transparent,  colorless,  and  as  far  as  can  be  determined 


THE   CELL   DOCTRINE.  77 

by  the  highest  powers,  structureless,  being  visible  only 
through  its  difference  in  refracting  power  as  com- 
pared with  the  menstruum  in  which  it  floats,  or  by 
the  granular  matter  it  may  entangle;  and  these 
characters  are  the  same  at  every  period  of  its  exist- 
ence. They  may  be  studied  in  the  simplest  vegetables, 
in  the  thallus  of  the  sugar  fungus,  among  the  low^est 
animals,  in  the  amoeba  (Plate,  Fig.  16),  and  in  higher 
animals  in  the  mucous,  pus,  or  white  blood  corpus- 
cles (Plate,  Fig.  10),  all  of  which  are  composed  almost 
purely  of  germinal  matter ;  the  very  thin  periphery 
of  formed  material  being  scarcely  appreciable  or  dis- 
tinguishable from  the  diffraction  band. 

In  its  endowments  and  properties,  germinal  matter 
is  actingy  living,  groiving,  and  moving,  through  some 
inherent  power  of  its  own.  It  alone,  as  stated,  is 
capable  of  producing  material  like  itself  out  of  pab- 
ulum, and  multiplying  by  division,  or  a  dropping  oft 
of  a  portion  of  itself,  which  portion  immediately 
assumes  an  independent  existence,  and  grows,  main- 
tains, and  reproduces  itself  like  the  parent  germinal 
matter.  It  is  also  capable  of  being  stained  by  an 
ammoniacal  solution  of  carmine,  and  the  younger  it 
is,  or  more  recently  formed,  the  deeper  is  the  stain  it 
assumes.  And  since  the  latest  formed  always  ap- 
pears in  the  centre  of  the  mass,  successive  tints,  or 
zones  of  color,  will  often  be  produced  in  the  staining 
process,  growing  deeper  from  without  inward,  as 
seen  in  Fig.  17  of  Plate. 

It  has  been  stated  that  what  is  called  nucleus  by 
Virchow  and  others,  is  included  in  germinal  matter. 
This  is  true,  though  the  nucleus  is  not  always  the 

7* 


78  THE   CELL   DOCTRINE. 

whole  of  the  germinal  matter.  There  maybe  other 
older  germinal  matter  beyond  the  nucleus,  on  its 
way  to  conversion  into  formed  material,  but  still 
germinal  matter,  which  assumes  a  tint  with  carmine, 
but  not  so  deep  as  the  nucleus.  Thus,  the  entire 
mass  of  the  pus  corpuscle  (Plate,  Fig.  10),  except  its 
extreme  periphery,  is  germinal  matter,  yet  there  is 
within  this  another  younger  portion  of  germinal 
matter,  taking  a  deeper  tint  with  carmine,  but  which 
alone  of  the  elements  of  this  cell,  we  are  in  the  habit 
of  calling  "  nucleus."  The  "  nucleus^''  then,  is  noth- 
ing but  a  new  centre  of  germinal  matter,  and  the 
^' nucleolus  ^^  is  a  younger  centre.  And  there  may 
even  be  within  this,  a  still  younger  portion  of  living 
matter,  taking  even  a  deeper  stain,  which  might  be 
called  a  ^'micleoleolusJ^  By  this  staining  process  may 
we  distinguish  the  nucleolus  from  a  minute  oil-drop 
often  mistaken  for  it,  and  which  will  not  assume  any 
tint. 

On  the  other  hand,  germinal  matter  in  a  compar- 
atively quiescent  state  is  often  quite  destitute  of 
nuclei.  But  let  the  mass  be  freely  supplied  with 
nutrient  matter  and  nuclei  and  nucleoli  rapidly 
make  their  appearance. 

So  with  regard  to  the  "cell  contents"  over  and 
above  the  nucleus,  although  they  may  all  be  germinal 
matter,  yet  this  is  not  necessarily  the  case.  Thus  in 
the  white  blood  corpuscle  and  mucous  corpuscles, 
what  Yirchow  would  consider  cell  contents  is  all  ger- 
minal matter;  but  the  superficial  epithelial  cell  lining 
the  interior  of  the  mouth  has  its  nucleus  alone  com- 
posed of  germinal  matter,  and  much  that  has  been  de- 


THE   CELL   DOCTRINE.  79 

scribed  as  cell  contents  is  really  formed  matter.  (Figs. 
5  and  6  of  Plate.)  More  nearly  does  the  germinal  mat- 
ter of  Beale*  correspond  with  the  "protoplasm"  of 
Max  Schultze,  with  which,  indeed,  it  seems  identical, 
except  that  the  latter  observer  seems  somewhat  at  a 
loss  how  to  dispose  of  the  nucleus,  of  which  he  does 
not  speak  as  a  new  or  young  centre  of  protoplasm. 

Formed  material,  instead  of  being  aeiive,  so  far  as 
the  vital  acts  described  as  characteristic  of  germinal 
matter  are  concerned,  i^jpassive,  non-acting,  dead,  and 
can  only  increase  at  the  expense  and  death  of  the 
germinal  matter,  on  the  periphery  of  which  it  is 
formed.  It  differs  widely  in  its  appearance,  and  is 
often  "structured"  as  in  muscle  and  nerve,  but  not 
necessarily  so,  as  is  seen  in  the  intercellular  sub- 
stance of  hyaline  cartilage.  It  possesses  also  cer- 
tain properties,  different  in  different  situations,  and 
widely  different  also  from  those  of  germinal  matter. 
Thus  it  is  contractile  in  the  sarcous  tissue  of  muscle, 
exhibits  neurility  in  the  nerve,  is  protective  in  epi- 
thelium, is  diffluent  as  the  formed  material  of  the 
milk-cell  (milk),  and  in  the  formed  material  of  the 
liver-cell  (bile).  Again,  it  is  hard  and  elastic  in  the 
intercellular  substance  of  cartilage,  and  epidermis, 
horn  and  nails.  It  does  not  become  stained  on  being 
soaked  in  weak  solution  of  carmine  in  ammonia,  and 
if  by  reason  of  the  strength  of  the  solution  it  should 
happen  to  be  stained,  the  color  will  w^ash  out  on 
soaking  in  glycerine,  which  is  not  the  case  with  the 
coloring  of  the  germinal  matter. 

*  Beale,  Protoplasm;  or,  Life,  Force,  and  Matter.     London: 
1870,  p.  38. 


80  .  THE   CELL   DOCTRINE. 

The  cause  of  this  permanent  staining  of  the  ger- 
minal matter  by  an  ammoniacal  solution  of  carmine, 
is  thought  by  Dr.  Beale  to  be  due  to  an  acid  reaction 
of  this  matter,  in  consequence  of  which  the  carmine 
is  precipitated  from  its  alkaline  solution.  This  view 
would  seem  to  be  confirmed  by  the  researches  of 
Ranke  on  the  Reaction  of  the  Tissues. 

The  size  of  the  elementary  part,  as  thus  composed, 
is  extremely  various.  The  smallest  particles  of  ger- 
minal matter,  measured  by  Dr.  Beale,  are  less  than 
tuAtju  ^f  ^^  i^^<^^  i^^  diameter,  and  w^ould  not  be 
called  cells  in  the  ordinary  sense  of  the  word,  yet 
they  are  functionally  such ;  that  is,  they  grow,  multi- 
ply by  division,  and  under  appropriate  circumstances 
assume  the  characters  of  fully  formed  cells.  On  the 
other  hand,  the  largest  epithelial  cells,  including 
their  germinal  matter  and  formed  material,  are  often 
as  large  as  the  ^J^  of  an  inch  in  diameter,  or  larger; 
cells  of  morbid  growths  are  sometimes  ^l^,  while  the 
human  ovum,  which  is  a  typical  cell,  varies  from  the 
24U  t^  T  Ju  of  an  inch.  Pure  germinal  matter  is  rarely 
seen  in  masses  as  large  as  the  ^  J^  of  an  inch  in  di- 
ameter, without  breaking  up  into  smaller  particles  of 
germinal  matter,  and  as  constituting  the  nuclei  of 
fully  formed  cells,  i^  usually  from  g^'^^  to  -j^'^^  of  an 
inch  in  diameter. 

The  method  of  production  of  formed  material  is  best 
studied  in  the  epithelial  structures,  particularly  in 
the  epithelium  lining  mucous  cavities,  of  which  sec- 
tions may  be  easily  made  down  to  the  vessels  whence 
their  nourishment  is  obtained.  In  the  deep  layers, 
next  the  nutrient  surface,  the  cells  will  be  found  to 


THE    CELL   DOCTRINE.  81 

consist  of  almost  pure  germinal  matter  (Plate,  Fig. 
1),  imbedded  in  a  soft,  mucus-like,  yet  continuous 
formed  matter.  These  masses  of  germinal  matter 
divide  and  subdivide,  pushing  the  older  masses  to- 
wards the  surface,  to  make  up  for  those  which  are 
constantly  exfoliated.  While  this  is  going  on,  how- 
ever, the  germinal  matter  keeps  increasing  in  size 
until  the  cells  arrive  half  way  towards  the  surface,  by 
absorption  of  nutrient  pabulum,  which  has  to  work 
its  way  through  any  formed  material  already  exist- 
ing. At  the  same  time,  a  portion  of  the  germinal 
matter  is  being  converted  into  formed  material, 
which  accumulates  on  its  surface,  within  that  already 
formed,  stretching  it,  and  becoming  more  or  less  in- 
corporated with  it.  Thus,  both  constituents  of  the  cell 
increase  up  to  a  certain  point,  the  cells  constantly  growing 
in  consequence.  As  new  cells  are,  however,  produced 
from  below,  the  older  ones  are  removed  farther  and 
farther  away,  the  formed  matter  becoming  more  and 
more  impervious  to  nutrient  pabulum.  At  length  a 
point  is  attained  when  the  entire  cell  ceases  to  in- 
crease in  size,  since  no  pabulum  reaches  the  masses 
of  germinal  matter,  though  the  latter  is  still  being 
converted  into  formed  material.  Hence,  the  masses 
of  germinal  matter  actually  grow  smaller,  as  the  cell 
increases  in  age;  and  when  the  periphery  is  reached, 
there  remains  but  a  small  nucleus  of  germinal  matter, 
with  a  large  quantity  of  formed  material.  Thus,  we 
are  enabled  to  judge  of  the  age  of  the  cell  by  the 
relative  quantity  of  germinal  matter  and  formed  ma- 
terial ;  if  the  former  be  large,  and  the  latter  small, 
the  cell  is  young,  whereas,  if  the  opposite  relation 


82  THE   CELL   DOCTRINE. 

exists,  the  cell  is  old,  and  almost  ready  to  exfoliate. 
But  exfoliation  probably  does  not  take  place  until 
the  last  particle  of  germinal  matter  dies,  and  the  en- 
tire cell  becomes  a  mass  of  passive,  lifeless,  formed 
material. 

The  production  of  formed  material  from  germinal 
matter,  may  also  be  studied  in  the  conversion  of  the 
white  blood-corpuscle  into  the  red.  In  the  spring  of 
the  year,  many  white  corpuscles  can  be  found  in  the 
blood  of  tbe  frog  and  newt,  undergoing  conversion 
into  formed  material  at  their  edges,  where  the  usual 
granular  appearance  is  being  substituted  by  the 
smooth  and  slightly  colored.  This  goes  on  until  all 
except  the  nucleus  is  thus  converted.  In  mammalia 
this  change  goes  on  until  the  whole  white  corpuscle 
is  thus  converted  into  the  red. 

Secondary  Formed  Material,  —  There  are  certain' 
kinds  of  formed  material,  to  which  this  term  is  ap- 
plied by  Dr.  Beale.  These  are  the  oil  of  the  fat  cell 
or  vesicle,  and  the  starch  granule  of  the  vegetable 
cell.  It  results,  as  does  all  formed  material,  by  a  con- 
version of  the  germinal  matter  into  this  special  form. 
The  young  fat  cell,  as  all  young  cells,  is  almost  pure 
germinal  matter;  as  it  grows  older,  however,  and  is 
exposed  to  oxidizing  influences,  the  conversion  of 
germinal  matter  takes  place,  partly  into  the  cell  wall 
proper  of  the  fat  vesicle,  and  partly  into  the  second- 
ary formed  material  or  oil,  until  finally,  it  becomes  a 
mere  dot  on  the  inner  surface  of  the  cell-wall,  or  dis- 
appears altogether. 

The  increase  of  cells,  according  to  Beale,  takes  place 
in  several  ways ;  every  cell  coming  from  a  pre-existing 


THE    CELL    DOCTRINE.  83 

cell,  but  the  germinal  matter  is  always  the  portion 
in  which  it  originates. 

There  is  not  generally  a  symmetrical  division  of 
the  nucleus  into  two,  and  these  into  four,  as  is  so 
often  described,  and  as  is  often  seen  in  the  vegetable 
cell,  but. there  is  rather  a  budding,  and  subsequent 
dropping  ofi*  of  the  portions  of  germinal  matter 
which  is  to  produce  the  new  cell,  and  which  almost 
always  assumes  the  spherical  form  when  allowed  to 
float  freely.  (See  Figure  10  of  Plate.)  The  formed 
material  is  never  active,  but  entirely  passive  in  the 
process  of  cell  multiplication. 

Nutriiion  of  Cells. — So,  too,  in  the  nutrition  of  the 
cell,  the  germinal  matter  is  the  sole  active  agent. 
The  formed  material  may  act  as  a  filter  to  the  nu- 
trient matter,  but  is  quite  passive.  The  pabulum, 
which  is  coursing  through  the  bloodvessels,  becomes 
converted  into  germinal  matter,  which  in  turn  be- 
comes formed  material,  and  so  long  as  this  is  kept 
up,  the  cell  continues  to  grow.  The  course  taken  by 
the  pabulum,  and  the  order  of  conversion,  is  shown 
by  the  arrows,  in  Figure  17,  of  Plate,  and  will  be 
readily  understood  by  reference  to  the  explanation. 
Occasionally,  and  especially,  in  disease,  the  formed 
material  may  become  the  pabulum  for  rapidly  mul- 
tiplying cells,  and  thus  be  consumed. 

Intercellular  substance  has  already  been  spoken  of 
as  formed  material.  We  have  it  most  strikingly  pres- 
ent in  the  white  fibrous  tissue,  or  tissue  of  tendons, 
and  in  hyaline  cartilage.  If  the  former  be  stained 
by  carmine,  and  examined  in  thin  section  under  the 
microscope,  it  will  be  found  composed  of  beautiful 


84  THE   CELL   DOCTRINE. 

bands  of  gently  waving  fibrous  tissue,  or  tissue  ex- 
hibiting a  fibrous  appearance,  at  varying  intervals  in 
whicb  are  noted  nuclear  masses  of  germinal  matter, 
which  have  assumed  the  tint  of  carmine.  Or,  if  di- 
lute acetic  acid  be  added  to  the  specimen,  the  fibrous 
appearance  will  at  once  become  homogeneous,  while 
the  nuclei  will  be  rendered  distinct,  and  clearly  visi- 
ble. In  young  tendon  (Plate,  Figure  11),  the  masses 
of  germinal  matter  will  be  found  very  abundant,  and 
the  intercellular  fibrous  substance  in  smaller  quan- 
tity than  in  old  tendon  where  the  masses  of  germinal 
matter  occur  only  at  long  intervals.  These  masses  of 
germinal  matter,  or  connective  tissue  corpuscles,  it 
will  be  recollected,  are  considered  by  Yirchow  as 
perfect  cells,  presenting  cell  wall,  cell  contents,  and 
nucleus,  and  the  fibrous  intercellular  substance  as  a 
subsequent  modification  of  a  homogeneous  matrix, 
dropped  between  the  cell  by  the  bloodvessels.  These 
connective  tissue  corpuscles  are  regarded  by  Beale 
as  simple  masses  of  germinal  matter,  the  conversion 
of  which  into  formed  material  produces  the  fibrous 
intercellular  substance,  as  seen  in  Figure  11,  Plate, 
and  between  which  and  the  intercellular  substance 
immediately  adjoining,  there  is  no  line  of  separation, 
constituting  a  cell  wall. 

As  the  tendon  grows  older,  the  masses  of  germinal 
matter  become  less  abundant,  because  a  larger  num- 
ber have  been  totally  converted  into  formed  material; 
and  the  bands  of  indestructible  material  which  some- 
times join  them,  and  which  are  considered  by  Virchow 
as  a  part  of  his  canalicular  system,  are,  according  to 
Beale,  nothing  hut  imperfectly  converted  formed  material, 


THE   CELL   DOCTRINE.  85 

or  rather  germinal  matter,  which  has  not  been  con- 
verted. While  the  twisted  and  curling  cord-like 
fibres  of  the  so-called  yellow  elastic  tissue,  also  con- 
sidered by  Virchow  as  a  part  of  his  canalicular  sys- 
tem, are  thought  by  Beale  to  be  composed  in  part  of 
true  yellow  elastic  tissue,  such  as  is  found  in  the 
ligamentum  nuchge,  and  likewise  formed  from  nu- 
clei (Plate,  Figure  14),  but  in  part  also  of  the  remains 
of  nerves,  and  vessels,  which  were  active  at  an  earlier 
period  of  life.* 

So,  also,  with  hyaline  cartilage.  According  to  Beale, 
the  intercellular  substance  results  from  the  conver- 
sion of  the  so-called  cartilage  corpuscles  or  cells  into 
formed  material,  and  here  also  the  germinal  matter 
is  directly  continuous  with  the  matrix,  no  proper  cell 
wall  intervening. 

Cartilage  is  not  to  be  considered  as  a  distinct  class 
of  tissue  from  epithelium,  nor  can  the  latter,  in  all 
cases  be  distinguished  from  cartilage  by  the  exist- 
ence of  separate  cells,  since  in  many  forms  of  epi- 
thelium, at  an  early  period  of  existence,  the  formed 
material  corresponding  to  the  masses  of  germinal 
matter  is  continuous  throughout,  and  presents  no 
indication  of  division  into  cells.f  A  "  cell,"  or  ele- 
mentary part,  then,  of  fully  formed  tendon  or  carti- 
lage, would  consist  of  a  portion  of  germinal  matter, 
with  a  proportion  of  formed  material  about  it,  ex- 
tending to  a  line  midway  between  that  mass  of  ger- 

*  Beale,  On  the  Structure  and  Growth  of  the  Tissues,  and  on 
Life.     London :  1865,  pp.  95,  96,  and  101. 

f  Beale,  Protoplasm;  or,  Life,  Force  and  Matter.  London: 
1870,  p.  61. 


86  THE    CELL   DOCTRINE. 

minal  matter  and  the  masses  immediately  adjacent, 
of  which  the  .cartilage  or  tendon  is  composed ;  and 
such  a  line  would  correspond  to  the  outer  part  of 
the  surface  of  an  epithelial  cell.*  In  very  j^oung  car- 
tilages, as  in  very  young  epithelium,  the  cells  consist 
of  germinal  matter  only,  with  a  small  quantity  of 
soft  formed  material  intervening;  and  to  understand 
the  true  relation  of  the  cells  to  the  intercellular  sub- 
stance, the  tissue  should  be  studied  at  different  pe- 
riods of  its  growth. 

So,  too,  a  "  cell "  or  elementary  part  of  muscle  or 
nerve,  would  consist  of  a  mass  of  germinal  matter 
(the  so-called  nucleus),  with  a  portion  of  muscular 
or  nervous  tissue  corresponding  with  it,  and  with 
which  it  is  uninterruptedly  continuous. 

In  the  formation  of  the  contractile  tissue  of  muscle^ 
the  germinal  matter  seems  to  move  onward,  under- 
going conversion  at  its  posterior  part,  into  the  mus- 
cular tissue,  while  it  maintains  itself  by  absorbing 
and  converting  pabulum.  This  will  be  understood 
by  reference  to  Fig.  13.  The  fibres  of  yellow  elastic 
tissue  are  formed  in  precisely  the  same  manner.  (See 
Plate,  Fig.  14.)  Nerve  fibres,  which  in  their  com- 
pleted state  consist  almost  wholly  of  formed  material, 
are  similarly  produced.  In  the  young  state,  the  fibre 
is  composed  of  masses  of  germinal  matter,  linearly 
arranged,  and  in  close  proximit3^  As  the  conversion 
takes  place  and  the  fibre  is  produced,  these  become 
more  widely  separated,  and  the  tissue  resulting  from 
such  conversion  is  nerve.  (Plate,  Fig.  15.) 

*  Beale,  Protoplasm,  pp.  51-2. 


THE    CELL   DOCTRINE.  87 

The  '^Cell^'  or  ^'Elementary  Fari'^  in  Disease, 

Here,  as  in  normal  nutrition,  the  germinal  matter 
is  alone  active.  It  is  impossible  to  state  precisely 
every  instance,  but  it  is  probable  that  in  the  majority 
of  cases  of  disease,  the  morbid  state  consists  es- 
sentially in  a  modification  of  the  healthy  nutrition 
of  the  cell,  that  is,  the  cell  is  made  to  grow  more  or 
less  rapidly,  or  is  perverted  in  its  mode  of  growth, 
though  it  is  likely  that  within  certain  limits,  the  con- 
ditions under  which  cells  ordinarily  live  may  be 
modified  without  deviation  from  health.  But  in  in- 
flammatory  processes  attended  by  local  products,  as 
jpus  or  lymph,  and  in  the  production  of  tubercle  and 
cancer  we  see  the  results  of  excessive  multiplication 
and  perversion  of  germinal  matter  consequent  upon 
the  appropriation  of  an  excess  of  nutrient  'pabulum. 
In  other  instances,  as  cirrhosis,  where  there  is  shrink- 
ing, and  hardening,  and  wasting,  we  see  the  efl*ects 
of  a  diminished  supply  of  pabulum,  either  through 
a  diminution  in  the  quantity  supplied,  or  an  imper- 
meability in  the  septum  through  which  it  is  com- 
pelled to  pass. 

An  increased  supply  of  pabulum  may  be  admit- 
ted to  germinal  matter,  either  in  consequence  of 
the  removal  of  barriers  through  which  it  is  ordi- 
narily compelled  to  pass,  or  in  consequence  of  the 
nature  of  the  fluids  hy  which  it  is  bathed.  A  sim- 
ple illustration  is  seen  in  suppuration  in  epithelium, 
or  the  germinal  matter  of  any  tissue;  for,  according 
to  Beale,  suppuration  and  morbid  processes  gene- 
rally, are  not  restricted  to  any  one  kind  of  ger- 


88  THE   CELL   DOCTRINE. 

minal  matter,  as  the  connective  tissue  corpuscle,  but 
may  occur  in  all  germinal  matter  to  which  the  con- 
ditions are  supplied.  Using  epithelium  by  way  of 
illustration,  as  the  result  of  the  increased  supply  of 
pabulum,  the  germinal  matter  first  grows,  as  seen  in 
Plate,  Figs.  7  and  8,  then  in  the  luxuriance  of  its 
growth,  even  at  the  expense  of  the  formed  matter, 
sends  out  buds  or  processes,  which  soon  drop  ofi'and 
become  separate  pus  corpuscles.  (Figs.  9  and  10.) 
These  are  produced  so  rapidly  that  there  is  not  time 
for  formed  material  to  form  upon  their  surface  in  any 
quantity,  and  they  have  not. time,  therefore,  to  pass 
on  into  epithelium.  Hence  pus  corpuscles  are  al- 
most pure  germinal  matter.  So  soon  as  the  process 
ceases,  in  consequence  of  the  supply  of  pabulum 
being  diminished,  the  germinal  matter  multiplies 
less  rapidly;  opportunity^  is  permitted  for  the  pro- 
duction of  formed  material  on  its  periphery,  and  the 
cell  now  passes  through  the  different  grades  of  epithe- 
lium, as  described  on  pages  80,  81,  and  82.  The  pus 
corpuscles  are  analogous  to  the  deepest  layers  of 
epithelial  cells  there  referred  to,  which  deep  cells 
are  in  fact  the  "  mucous  corpuscles,"  so-called,  well 
known  to  be  morphologically  identical  with  pus  cor- 
puscles; the  former  being  simply  the  young  epithe- 
lial cell  on  its  w^ay  to  become  perfect  epithelium, 
while  the  latter  is  the  same  also,  though  never  al- 
lowed to  pass  into  the  perfectly  formed  state. 

Again,  in  piieumojiia,  and  here  we  note  where  the 
paths  of  Yirchow  and  Beale  separate  more  widely, 
the  so-called  "  exudation,"  or  product  which  fills 
up  the  vesicular  portion  of  the  lung,  considered  by 


THE   CELL   DOCTRINE.  89 

Vircliow  a  local  one,  the  result  of  a  proliferation  of 
the  connective  tissue  corpuscle  of  the  part,  and 
having  no  dependence  upon  the  blood,  is  regarded 
by  Beale  as  the  result  of  a  proliferation  of  minute 
'particles  of  (jerminal  matter  (very  much  smaller  than 
white  blood  corpuscles),  which  have  passed  out 
through  the  capillary  walls  with  the  liquor  sanguinis. 

•  In  all  inflammatory  processes  and  fevers,  this  is 
believed  by  Dr.  Beale  to  take  place  to  a  greater  or 
less  extent,  the  little  masses  of  germinal  matter  or 
nuclei  in  the  capillary  walls  also  taking  part,  often 
increasing  in  size  to  such  degree  that  they  materially 
obstruct  the  passage  of  the  blood,  and  by  dropping 
oft*  portions  give  rise  to  bodies  floating  in  the  blood 
precisely  similar  to  white  blood  corpuscles,  or  pus 
corpuscles;  indeed.  Dr.  Beale  considers  that  this  may 
be  one  of  the  sources  of  origin  of  the  white  blood  cor- 
puscle.* 

So,  also,  tubercle  is  believed  by  Dr.  Beale  to  result 
either  from  the  multiplication  of  masses  of  germinal 
matter  which  have  passed  through  the  capillary  walls 
from  the  blood,  or  from  the  masses  of  germinal  matter 
usually  termed  nuclei,  in  connection  with  the  capillary 
walls.  He  says,  in  illustration,  "In  a  case  of  tubercle, 
which  was  very  rapidly  developed  upon  the  surface 
of  the  pia  mater,  in  a  man  of  tubercular  constitu- 
tion, I  proved  most  distinctly,  that  the  tubercles  were 
connected  with  the  vascular  walls,  and  that  if  the 
nuclei  had  not  given  origin  to  them,  they  were  cer- 

*  Beale,  Microscope  in  Clinical  Medicine,  3d  ed.  London  : 
1867,  p.  166. 

8* 


90  THE    CELL   DOCTRINE. 

tainly  implicated.  My  own  opinion  is,  that  these 
nuclei  gave  origin  to  the  tubercle  corpuscles,  in  con- 
sequence of  receiving  from  the  blood  peculiar  nu- 
trient matter.  In  the  lung  I  have  seen  appearances 
which  point  to  a  similar  conclusion."*  Would  not 
these  views  arise  from  appearances  precisely  analo- 
gous to  those  represented  as  giving  support  to  the 
view,  that  tubercle  originates  in  the  perivascular 
sheaths  of  bloodvessels  ?  The  views  of  Beale,  II. 
Charlton  Bastian,t  and  Cornil,J  would  then  consti- 
tute simply  diflerent  modes  of  expression  of  the  same 
truths. 

ROBIN,  E.  A.,  POUCHET,§  1867. 

Robin,  who  may  be  considered  the  mouthpiece  of 
the  French  school  of  histologists,  reduces  the  human 
body  to  elementary  parts,  usually  microscopic,  which 
he  calls  anatomical  elements.  The  forms  he  makes 
threefold,— ^6re5,  tubes,  and  cells. 

The  fibres  are  generally  of  considerable  length, 
sometimes  extending  from  the  lower  part  of  the 

*  Microscope  in  Clinical  Medicine,  3d  ed.,  1867,  p.  205. 

f  Bastian,  H.  C,  Tuberc.  Meningitis.  Edinb.  Med.  Jour.,  1867, 
p.  875. 

%  Cornil,  Tubercle  in  Connection  with  the  Vessels.  Archiv.  do 
Phys.  Norm,  et  Path.,  Jan.  et  Pev.,  1868. 

§  Our  information  with  regard  to  M.  Eobin's  views,  is  de- 
rived from  an  admirable  exposition  of  them  published  in  vol.  iv, 
1867,  of  the  New  York  Medical  Journal,  by  Dr.  Wm.  T.  Lusk, 
who  there  states  that  he  has  them  mainly  from  a  course  of  familiar 
and  private  instruction,  furnished  to  him  by  M.  C.  H.  Georges 
Pouchet,  son  of  the  eminent  physiologist,  Prof.  F.  A.  Pouchet, 
Assistant  to  M.  Kobin,  Lecturer  upon  Anatomy  and  Histology  to 
the  Ecole  Pratique,  author  of  "Un  Precis  d'Histologie,"  &c. ;  so 
that  they  may  be  said  to  be  the  views  also  of  the  elder  Pouchet. 


THE   CELL   DOCTRINE.  91 

spinal  cord  to  the  extremity  of  the  foot.  Their  di- 
ameter is,  however,  small,  often  not  exceeding  .001 
millimetre,  or  .00003987  of  an  inch. 

The  tubes  offer  as  objects  of  study  the  walls  and 
the  cavity. 

The  cells  of  vegetables  have  a  wall,  a  cavity,  and 
contents  (air,  oil,  &c.).  The  cells  of  animals,  on  the 
contrary,  are,  as  a  rule,  homogeneous.  Animal  cells 
containing  a  cavity  are  only  found  exceptionally.  The 
substance  of  cells  is  ordinarily  granular.  Most  cells 
contain  an  ovoid  nucleus  more  granular  than  the 
substance  itself. 

In  all  cells  the  nuclei  afford  different  chemical  re- 
actions from  those  of  the  substance  of  the  element. 
Each  cell  is  an  independent  organism,  passing  through 
various  stages  of  development,  from  birth  to  death. 

The  birth  (origin)  of  the  elements  takes  place  by 
1st,  segmentation;  2d,  genesis;  3d,  epigenesis;  4th, 
germination. 

1st.  Segmentation. — The  human  ovum  is  a  small 
hollow  sphere,  containing  in  its  interior  the  vitellus 
or  yolk,  which  consists  of  granular  matter  in  a  hya- 
line substance.  At  the  end  of  a  certain  time  parti- 
cles of  the  granular  matter  approximate,  unite  and  form 
a  nucleus  in  the  vitellus.  Next,  the  nucleus  elongates, 
takes  an  hour-glass  form  (biscuit),  then  divides.  The 
division  of  the  yolk  occurs  simultaneously.  In  the 
same  way,  the  division  takes  place  into  4,  8,  16,  and 
more  parts.  These  divisions  of  the  vitellus  have  re- 
ceived the  name  vitelline  globules.  Their  mode  of 
formation  is  called  segmentation. 

2d.   Genesis. — When  the  vitelline  globes  have  be- 


92  THE   CELL   DOCTRINE. 

come  very  small  by  successive  segmentation  (diame- 
ter .008  millimetre,  .00031396  of  an  inch),  these  little 
bodies  take  the  name  of  eriihryonic  cells. 

According  to  M.  Robin,  these  cells  dissolve. 
From  the  fusion  results  a  blastema,  in  the  midst  of 
which  nuclei  make  their  appearance.  This  is  known 
as  genesis.  It  is  the  second  and  most  frequent  mode 
of  the  formation  of  anatomical  elements.  It  is  char- 
acterized by  the  appearance  of  an  anatomical  ele- 
ment in  a  fluid  termed  blastema,  in  which  the 
element  did  not  previously  exist. 

3d.  Epigenesis. — When  the  embryonic  cells  dis- 
solve, the  embryo-plastic  nuclei  are  produced  by 
genesis  in  the  blastema  which  results  from  their 
fusion.  Then  little  cone-like  prolongations  of  trans- 
parent matter  are  observed  at  the  extremities  of  the 
nuclei,  giving  rise  to  the  fusiform  bodies,  which  are 
the  connective  tissue  corpuscles.  This  mode  of  for- 
mation by  growth  upon  another  element  is  known 
as  epigenesis,  and  is  the  mode  in  which  connective 
tissue  is  developed.  The  prolongations  of  these  fu- 
siform bodies  constitute  the  non-elastic  fibres  or 
white  fibrous  tissue  element  of  connective  tissue. 
Sometimes  the  substance  deposited  by  epigenesis 
upon  the  nucleus  has  several  prolongations,  forming 
a  stellate  cell  or  connective  tissue  corpuscle.  These 
fusiform  and  stellate  cells  are  likewise  known  as 
embryo-plastic  or  fibro-plastic  bodies,  and  this  latter 
term  is  a  most  common  one  in  French  histology. 

The  elastic  fibres  of  connective  tissue  are  likewise 
formed  by  epigenesis,  but  upon  special  nuclei,  and 
the  prolongations  are  insoluble  in  acetic  acid. 


THE    CELL    DOCTRINE.  93 

There  is  an  early  period  of  foetal  life,  previous  to 
the  formation  of  connective  tissue,  in  which  we  find 
only  embryo-plastic  nuclei  and  fusiform  bodies  in 
amorphous  matter.  This  is  called  embryo-plastic 
tissue.  Growth  at  this  epoch  is  most  rapid,  the 
foetus  reaching  in  a  short  space  of  time  the  dimen- 
sion of  .030  millimetre  (.0118  of  an  inch). 

4th.  Germination. — This  is  very  frequent  in  vege- 
tables, but  in  animals  only  one  example  is  known, 
viz.,  at  a  period  previous  to  the  fecundation  of  the 
ovum.  Before  segmentation  takes  place  the  vitellus 
is  observed  to  retract.  The  hyaline  substance  pushes 
out  a  prolongation,  which  becomes  round,  separates, 
and  constitutes  an  independent  anatomical  element 
exterior  to  the  vitellus,  and  bearing  no  part  in  the 
future  development  of  the  ovum. 

The  following  account  of  certain  special  elements 
illustrates  and  further  explains  the  views  of  M. 
Eobin.  Bed  blood  globules  (hematics),  diameter,  .007 
millimetre  (^-i-Q-Q  of  an  inch);  thickness  .002  millime- 
tre ( 12*55  of  an  inch).  Blood  globules  are  elastic, — a 
property  enabling  them  to  elongate,  and  pass  through 
capillaries  which  have  a  calibre  less  than  the  diam- 
eter of  the  blood  globule.  They  are  homogeneous 
throughout — i.  e,,  have  no  cell  wall.  Blood  globules  are 
formed  by  genesis  in  the  blood  plasma.  In  the  foetus 
they  make  their  appearance  before  the  white  blood 
globules  (leucocytes).  In  man  there  are  two  kinds 
of  red  blood  globules,  viz. :  fi.rst,  embryonic;  second, 
normal.  The  embryonic  blood  globules  are  double 
the  size  of  the  normal  ones.  They  have  a  slightly 
granular  nucleus,  situated  nearly  in  the  centre,  which 


94  THE   CELL   DOCTRINE. 

is  insoluble  in  acetic  acid.  The  normal  blood  glob- 
ules are  not  a  transformation  of  the  embryonic. 
They  appear  by  genesis  in  the  midst  of  the  blastema 
of  the  blood.  After  the  fourth  month,  the  embry- 
onic globules  cease  to  form,  and  as  the  mass  of  the 
blood  increases,  the  proportionate  number  diminishes 
with  great  rapidity. 

Leucocytes,  or  white  blood  globules,  are  found  in 
many  tissues,  in  the  blood,  on  the  surface  of  mucous 
membranes;  in  a  word,  they  are  the  pus  corpuscles.  In 
form,  they  are  round,  with  pale,  well-defined  borders, 
and  contain  extremely  fine  gray  granules.  They 
possess  a  very  thin  envelope,  and  a  granular  cell 
contents.  The  normal  diameter  is  .008  millimetre 
Guuu  ^^  ^^  inch).  On  the  addition  of  water,  the 
leucocytes  swell,  the  granular  particles  are  agitated 
by  a  peculiar  movement  (first  observed  by  Brown), 
and  finally,  a  considerable  number  of  these  particles 
unite,  so  as  to  form  two  or  three  little  masses,  that  have 
been  mistaken  for  nuclei.  Upon  the  addition  of  acetic 
acid  the  same  reaction  follows,  but  with  greater 
rapidity. 

The  mode  of  production  may  be  followed,  step  by 
step,  upon  the  surface  of  wounds,  especially  little 
ones.  At  first  a  hyaline  liquid  appears.  At  the  end 
of  a  couple  of  hours,  this  liquid  becomes  finely 
granular,  and  then  all  at  once,  in  the  midst  of  the 
granulations,  we  perceive  small  granular  bodies  an- 
alogous to  leucocytes,  ofiering  the  same  chemical  re- 
actions, but  measuring  only  .003  millimetre  (.000118 
of  an  inch)  in  diameter.  They  are,  in  fact,  leuco- 
cytes of  young  growth.     When  leucocytes  are  re- 


THE   CELL   DOCTRINE.  95 

tained  in  the  economy,  as  in  shut  sacs,  they  increase 
in  size,  and  reach  a  diameter  of  .012  millimetre 
(s^uTj  ^f  ^^^  inch).  Then  they  fill  with  fat  gran- 
ules, and  are  known  as  corpuscles  of  inflammation 
(exudation  corpuscles,  compound  granule  cells). 
Finally  the  substance  and  investing  membrane  of  the 
leucocytes  disappear,  the  granules  dissolve  and  are 
reabsorbed. 

Capillaries. — The  finest  capillaries  are  anatom- 
ical elements  of  tubular  form,  with  transparent  re- 
sistant walls  which  measure  .001  mm.  (.00003937  of 
an  inch)  in  diameter.  These  walls  contain  granular 
ovoid  nuclei,  which  project,  sometimes  exteriorly, 
sometimes  upon  the  inner  surface  of  the  tubes. 
These  nuclei  measure  .006  mm.  (.00023622  of  an 
inch)  in  the  transverse,  and  .008  mm.  (.00031596  of 
an  inch)  in  the  long  diameter.  Their  long  axis  is 
parallel  to  that  of  the  vessel.  The  finest  capillaries 
have  a  diameter  of  .007  mm.  (.0003756  of  an  inch), 
leaving  a  calibre  (after  deducting  the  walls),  of  .005 
mm.  (.00019685  of  an  inch),  or  .002  mm.  (.00007874 
of  an  inch)  less  than  the  average  diameter  of  the 
blood  globules  which  traverse  them. 

They  are  formed  as  follows :  1st,  In  new  tissues, 
hollow  projections  push  out  from  contiguous  cap- 
illaries, which  meet  and  unite  together.  2d,  A 
solid  filament  forms,  in  which  nuclei  make  their  ap- 
pearance. Subsequently,  the  filament  becomes  hol- 
low, and  its  nuclei  remain  the  nuclei  of  the  capillary. 

A  single  perusal  of  these  views  as  thus  illustrated, 
will  convince  the  reader  that  spontaneous  formation  is 
the  prevailing  .mode  of  origin  of  the  elements  of 


96  THE   CELL   DOCTRINE. 

tissues,  according  to  the  French  school.  Such  pe- 
rusal cannot  fail  to  convince  the  reader  also  of  the 
accuracy  of  description  of  the  fully  formed  elements 
described  by  Robin. 

PROF.    HUXLEY,*  1869. 

There  is  one  kind  of  matter  which  is  common  to 
all  living  beings,  and  that  matter  is  "^^ro/opZa^m," 
the  scientific  name  for  "the  physical  basis  of  life." 
In  illustration  from  vegetable  life,  each  stinging 
needle  or  hair  of  the  common  nettle  consists  of  a 
very  delicate  outer  case  of  wood,  closely  applied  to 
the  inner  surface  of  which  is  a  layer  of  semi-fluid  mat- 
ter, full  of  innumerable  granules  of  extreme  minute- 
ness. This  semijhdd  lining  is  protoplasm,  which  thus 
constitutes  a  kind  of  bag,  full  of  a  limpid  fluid,  and 
roughly  corresponding  in  form  with  the  interior  of 
the  hair  which  it  fills.  When  viewed  w^ith  a  sufli- 
ciently  high  magnifying  power,  the  protoplasmic 
layer  of  the  nettle  hair  is  seen  to  be  in  a  condition 
of  unceasing  activity.  Local  contractions  of  the 
whole  thickness  of  its  substance  pass  slowly  and 
gradually  from  point  to  point,  and  give  rise  to  the 
appearance  of  progressive  weaves,  just  as  the  bending 
of  successive  stalks  of  grain  \by  a  breeze  produces 
the  apparent  billows  of  a  grain-field. 

But  in  addition  to  these  movements,  and  inde- 
pendently of  them,  the  granules  are  driven,  in  rela- 
tively rapid  streams,  through  channels  in  the  proto- 

*  Protoplasm ;  or,  The  Physical  Basis  of  Life.  A  Lecture  by 
Prof.  Huxley,  delivered  in  Edinburgh,  Nov.  18th,  1868. 


THE   CELL   DOCTHINE.  97 

plasm  which  seem  to  have  a  considerable  amount  of 
persistence.  The  currents  in  adjacent  parts  com- 
monly take  similar  directions,  coursing  in  a  general 
stream  up  one  side  of  the  hair  and  down  the  other, 
though  partial  currents  also  exist  which  take  differ- 
ent routes;  so  that  sometimes  trains  of  granules 
.may  be  seen  coursing  swiftly  in  opposite  directions, 
within  a  twenty-thousandth  of  an  inch  of  each  other; 
and  occasionally  opposite  streams  come  in  direct 
collision,  and  after  a  longer  or  shorter  struggle,  one 
predominates.  The  cause  of  these  currents  seems 
to  lie  in  contractions  of  the  protoplasm  which  bounds 
the  channels  in  which  they  flow,  but  which  are  so 
minute  that  the  best  microscopes  show  only  their 
effects,  and  not  themselves. 

Among  the  lower  plants,  it  is  the  rule  rather  than 
the  exception,  that  contractility  should  be  still  more 
openly  manifested  at  some  periods  of  their  existence. 
The  protoplasm  of  Algse  and  Fungi  becomes,  under 
many  circumstances,  partially  or  completely  freed 
from  its  woody  case,  and  exhibits  movements  of  its 
whole  mass,  or  is  propelled  by  the  contractility  of 
one  or  more  vibratile  cilia. 

In  illustration  of  animal  protoplasm.  Prof  Hux- 
ley adduces  the  colorless  corpuscles  of  the  blood; 
which,  under  the  microscope,  at  the  temperature  of 
the  body,  exhibit  a  marvellous  activity,  changing 
their  forms  with  great  rapidity,  drawing  in  and 
thrusting  out  prolongations  of  their  substance;  and 
creeping  about  as  if  they  wxre  independent  organ- 
isms. "  The  substance  which  is  thus  active  is  a 
mass  of  protoplasm,  and  its  activity  differs  in  detail 

9 


98  THE   CELL   DOCTRINE. 

rather  than  in  principle  from  that  of  the  protoplasm 
of  the  nettle.  Under  sundry  circumstances  the  cor- 
puscle dies,  and  becomes  distended  into  a  round 
mass,  in  the  midst  of  which  is  seen  a  smaller  spheri- 
cal body,  which  existed,  but  was  more  or  less  hid- 
den, in  the  living  corpuscle,  and  is  called  its  nucleus. 
Corpuscles  of  essentially  similar  structure  are  to 
be  found  in  the  skin,  in  the  lining  of  the  mouth, 
and  scattered  through  the  whole  framework  of  the 
body.  I^ay,  more;  in  the  earliest  condition  of  the 
human  organism,  in  that  state  in  which  it  has  just 
become  distinguishable  from  the  egg  in  which  it 
arises,  it  is  nothing  but  an  aggregation  of  such  cor- 
puscles, and  every  organ  of  the  body  was,  once,  no 
more  than  such  an  aggregation.  Thus  a  nucleated 
mass  of  protoplasm  turns  out  to  be  what  may  be  termed 
the  structural  unit  of  the  human  body.  As  a  matter  of 
fact,  the  body,  in  its  earliest  state,  is  a  mere  mul- 
tiple of  such  units;  and,  in  its  perfect  condition,  it 
is  a  multiple  of  such  units,  variously  modified." 
The  formula  which  expresses  the  essential  structural 
character  of  the  highest  animal,  very  nearly  covers 
all  the  rest,  as  the  statement  of  its  powers  and 
faculties  covered  that  of  all  others.  "Beast  and 
fowl,  reptile  and  fish,  mollusk,  worm,  and  polype, 
are  all  composed  of  structural  units  of  the  same 
character,  namely,  masses  of  protoplasm  with  a  nu- 
cleus. There  are  sundry  very  low  animals,  each  of 
which,  structurally,  is  a  mere  colorless  blood  cor- 
puscle, leading  an  independent  life.  But,  at  the  very 
bottom  of  the  animal  scale,  even  this  simplicity  be- 


THE   CELL   DOCTRINE.  99 

comes  simplified,  and  all  the  phenomena  of  life  are 
manifested  by  a  particle  of  protoplasm  without  a  nucleus, 

"  What  has  been  said  of  the  animal  world  is  no  less 
true  of  plants.  Imbedded  in  the  protoplasm  at  the 
broad,  or  attached  end  of  the  nettle  hair,  there  lies 
a  spheroidal  nucleus.  Careful  examination  further 
proves  that  the  whole  substance  of  the  nettle  is  made 
up  of  a  repetition  of  such  masses  of  nucleated  proto- 
plasm, each  contained  in  a  wooden  case,  which  is 
modified  in  form,  sometimes  into  a  woody  fibre, 
sometimes  into  a  duct  or  spiral  vessel,  sometimes 
into  a  pollen  grain,  or  an  ovule.  Traced  back  to  its 
earliest  state,  the  nettle  arises  as  the  man  does,  in  a 
particle  of  nucleated  protoplasm.  And  in  the  low^est 
plants,  as  in  the  lowest  animals,  a  single  mass  of  such 
protoplasm  may  constitute  the  whole  plant,  or  the 
protoplasm  may  exist  without  a  nucleus.  Under 
these  circumstances  it  may  well  be  asked,  how  is  one 
mass  of  non-nucleated  protoplasm  to  be  distinguished 
from  another?  why  call  one  *  plant,'  and  the  other 
*  animal?'  The  only  reply  is  that,  so  far  as  form  is 
concerned,  plants  and  animals  are  not  separable,  and 
that,  in  many  cases,  it  is  a  mere  matter  of  conven- 
tion whether  we  call  a  given  organism  an  animal  or 
a  plant." 

The  researches  of  the  chemist  have  also  shown  a 
like  uniformity  of  chemical  composition  in  "  proto- 
plasm "  or  living  matter,  proving  that  whatever  its 
source,  it  contains  carbon,  hydrogen,  oxygen,  and 
nitrogen,  producing  in  their  combination  a  complex 
substance,  whiib  iH»q\i)^  i'gi:^i>ai;ce^of  jtf  mor^;6xact 
nature,  we  Q2i\\'pfoieinaceousdv  Ulbuvlirioid  muittr. 


100  THE   CELL   DOCTRINE. 

Thus  far  it  is  plain  that  the  views  of  Prof.  Huxley 
accord  with  those  of  many  eminent  histologists  and 
physiologists,  the  result  of  whose  observations  have 
been  embodied  in  these  pages,  and  his  descriptions 
will  be  accepted  as  undoubtedl}^  accurate.  More 
widely,  in  common  with  the  school  of  so-called 
"  physicists,"  of  which  he  is  one,  does  he  differ  in 
his  views  in  that  which  is  yet  to  be  considered,  the 
origin  and  ultimate  fate  of  this  "protoplasm,"  or 
matter  of  life.  According  to  Prof.  Huxley,  the 
matter  of  life  is  composed  of  ordinary  matter,  and 
again  resolved  into  ordinary  matter  when  its  work 
is  done.  Waste  is  constantly  going  on  which  must 
be  supplied  by  food,  which  is  converted  into  proto- 
plasm. A  solution  of  smelling  salts  in  water,  with 
an  infinitesimal  proportion  of  some  other  saline  mat- 
ters, contains  all  the  elementary  bodies  which  enter 
into  protoplasm,  yet  an  animal  cannot  make  proto- 
plasm. And  this  is  characteristic.  It  must  take  it 
ready  made  from  some  other  animal  or  some  plant, 
the  animal's  highest  feat  of  constructive  chemistry 
being  to  convert  dead  protoplasm  into  the  living 
matter  of  life,  which  is  appropriate  to  itself  There- 
fore, in  seeking  for  the  origin  of  protoplasm,  we 
must  eventually  turn  to  the  vegetable  world.  The 
plant,  however,  takes  carbonic  acid,  water,  and  am- 
monia, and  converts  it  to  the  same  stage  of  living 
protoplasm  with  itself,  though  some  of  the  fungi 
need  higher  compounds  to  start  with;  and  no  plant 
can  live  on  the  uncompounded  elenients  of  proto- 
plasni^^djthe  abseilce  of  aay.^Re'^t  the  elements 
renders  the  ptant  unable  to  Tnariufacture  protoplasm. 


THE   CELL   DOCTRINE.  101 

These  elements,  carbon,  hydrogen,  ox3^gen,  and  ni- 
trogen, are  related  to  the  protoplasm  of  the  plant 
as  the  protoplasm  of  the  plant  to  the  animal.  But 
protoplasm  once  i^roduced,  all  the  phenomena  exhibited 
by  it  are  simply  its  properties,  just  as  the  phenomena 
exhibited  by  water  in  its  various  states  are  properties. 
They  do  not  take  place  through  the  guidance  of  any 
principle  called  "  vitality,"  any  more  than  the  phe- 
nomena of  water  take  place  by  virtue  of  "  aquosity." 
Prof.  Huxley  can  discover  no  halting-place  between 
the  admission  that  protoplasm  of  one  animal  or  vege- 
table is  essentially  identical  with  and  readily  con- 
verted into  another,  and  the  further  concession  that 
all  vital  action  may,  with  equal  propriety,  be  said  to 
be  the  result  of  the  molecidar  forces  of  the  protoplasm 
which  displays  it.  The  thoughts  to  which  we  give  utter- 
ance are  the  expression  of  molecular  changes  in  proto- 
plasm. These  are  admittedly  so-called  materialistic 
terms.  Yet  Prof.  Huxley  says :  "  I^fevertheless,  two 
things  are  certain:  the  one  that  I  hold  the  state- 
ment (above)  to  be  substantially  correct;  the  other, 
that  I,  individually,  am  no  materialist,  but  on 
the  contrary  believe  materialism  to  involve  grave 
philosophical  errors."  Such  union  of  materialistic 
terminology  with  the  repudiation  of  materialistic 
philosophy,  he  believes  to  be  "  not  only  consistent 
with,  but  necessitated  by  sound  logic."  This  he 
proceeds  to  show  in  this  manner :  If  it  be  supposed 
that  knowledge  is  absolute,  that  we  know  more  ot 
cause  and  eftect  than  a  certain  definite  order  of  suc- 
cession of  facts,  and  that  we  have  a  knowledge  of 
the  necessity  of  that  succession,  then  there  is  no 

9* 


102  THE    CELL   DOCTRINE. 

escape  from  utter  materialism  and  necessarianism. 
But  it  is  impossible  to  prove  that  anything  what- 
ever may  not  be  the  eflect  of  a  material  and  neces- 
sary cause,  and  no  act  is  really  spontaneous,  since  a 
really  spontaneous  act  is  one  which  has  no  cause. 
Yet  any  one  familiar  with  the  history  of  science  will 
admit  that  its  object  has  always  meant,  and  means 
the  extension  of  the  province  of  matter  and  causa- 
tion, and  the  concomitant  gradual  banishment  from 
all  regions  of  human  thought,  of  what  we  call  spirit 
and  spontaneity, — that  is,  the  object  of  all  science 
has  been  and  is  to  find  out  the  causes  of  all  phe- 
nomena; and  there  is  no  difference  between  the 
conception  of  life  as  the  product  of  a  certain  dispo- 
sition of  material  molecules  and  the  old  notion  of  an 
Archseus  governing  and  directing  blind  matter  within 
each  living  body,  except  that  here,  as  elsewhere,  mat- 
ter and  law  have  devoured  spirit  and  spontaneity. 
And  moreover,  the  physiology  of  the  future  will 
gradually  so  extend  the  realm  of  matter  and  law, 
until  it  is  coextensive  with  knowledge,  with  feeling, 
and  with  action.  It  is  this  j^rogress  of  knowledge^ 
according  to  Prof.  Huxley,  which  so  many  of  the 
best  minds  conceive  to  be  the  progress  of  material- 
ism^ which  they  watch  with  such  fear  and  powerless 
anger  as  a  savage  feels,  when,  during  an  eclipse,  the 
great  shadow  creeps  over  the  face  of  the  sun.  We 
know  nothing  of  this  terrible  *^  matter,"  except  as 
the  name  for  the  unknown  and  hypothetical  cause  of 
states  of  our  own  consciousness,  and  as  little  of  that 
"  spirit,"  except  that  it  is  also  a  name  for  an  un- 
known and  hypothetical  cause  of  states  of  conscious- 


THE   CELL   DOCTRINE.  103 

ness,  that  is,  matter  and  spirit  are  both  names  for 
the  imaginary  substrata  of  groups  of  natural  phe- 
nomena. Dire  necessity  and  "  iron  "  law  are  gratu- 
itously invented  bugbears.  If  there  be  an  "  iron  " 
law,  it  is  that  of  gravitation,  and  if  there  be  a  phys- 
ical necessity  it  is  that  a  stone  unsupported  will  fall 
to  the  ground.  We  know  nothing  more  of  this  latter 
phenomenon,  except  that  stones  always  have  fallen 
to  the  ground  under  these  conditions,  and  that  they 
will  continue  to  fall  to  the  ground  thus  unsupported. 

It  is  simply  convenient  to  indicate  that  all  the  con- 
ditions of  belief  in  this  case  have  been  fulfilled,  by 
calling  the  statement  that  unsupported  stones  will 
fall  to  the  earth  a  "  law  of  nature."  But  when  for 
will  we  exchange  must,  we  introduce  an  idea  of  neces- 
sity which  does  not  lie  in  the  observed  facts,  and  is 
not  warranted  by  anything  that  is  discovered  else- 
where. And  with  regard  to  which  Prof  Huxley 
says:  "For  my  part,  I  utterly  repudiate  and  anathe- 
matize the  intruder.  Fact  I  know,  and  Law  I 
know;  but  what  is  this  IlTecessity,  save  an  empty 
shadow  of  my  own  mind's  throwing?  But,  if  it  is 
certain  that  we  can  have  no  knowledge  of  the  nature 
of  either  matter  or  spirit,  and  that  the  notion  of 
necessity  is  something  illegitimately  thrust  into  the 
perfectly  legitimate  conception  of  law,  the  materialis- 
tic 'position  that  there  is  nothing  in  the  world  hut  matter, 
force,  and  necessity,  is  as  utterly  devoid  of  justification  as 
the  most  baseless  of  theological  dogmas. 

"  The  fundamental  doctrines  of  materialism,  like 
those  of  spiritualism,  and  most  other  *isms,'  lie  out- 
side *the  limits  of  philosophical  inquiry,'  and  David 


104  THE    CELL   DOCTRINE. 

Hume's  great  service  to  humanity  is  his  irrefragable 
demonstration  of  what  these  limits  are.  Hume 
called  himself  a  skeptic,  and  therefore  others  cannot 
be  blamed  if  they  apply  the  same  title  to  him;  but 
that  does  not  alter  the  fact  that  the  name,  with  its 
existing  implications,  does  him  gross  injustice.  If 
a  man  asks  me  what  the  politics  of  the  inhabitants 
of  the  moon  are,  and  I  reply  that  I  do  not  know; 
that  neither  I,  nor  any  one  else  have  any  means  of 
knowing;  and  that,  under  these  circumstances  I  de- 
cline to  trouble  myself  about  the  subject  at  all,  I  do 
not  think  he  has  any  right  to  call  me  a  skeptic.  On 
the  contrary,  in  replying  thus,  I  conceive  that  I  am 
simply  honest  and  truthful,  and  show  a  proper  regard 
for  the  economy  of  time.  So  Hume's  strong  and 
subtle  intellect  takes  up  a  great  many  problems 
about  which  we  are  naturally  curious,  and  shows  us 
that  they  are  essentially  questions  of  lunar  politics, 
in  their  essence  incapable  of  being  answered,  and 
therefore  not  worth  the  attention  of  men  who  have 
work  to  do  in  the  world."     .... 

"If  we  find  that  the  ascertainment  of  the  order  of 
nature  is  facilitated  by  using  one  terminology,  or  one 
set  of  symbols,  rather  than  another,  it  is  our  clear 
duty  to  use  the  former,  and  no  harm  can  accrue  so 
long  as  we  bear  in  mind  that  we  are  dealing  merely 
with  terms  and  symbols.  In  itself  it  is  of  little 
moment  whether  we  express  the  phenomena  of  mat- 
ter in  terms  of  spirit,  or  the  phenomena  of  spirit  in 
terms  of  matter ;  matter  may  be  regarded  as  a  form 
of  thought,  thought  may  be  regarded  as  a  property 
of  matter — each  statement    has  a  certain  relative 


THE    CELL    DOCTRINE.  105 

truth.  But  with  a  view  to  the  progress  of  science, 
the  materialistic  terminology  is  in  every  way  to  be 
preferred.  For  it  connects  thought  with  the  other 
phenomena  of  the  universe,  and  suggests  inquiry 
into  the  nature  of  those  physical  conditions,  or  con- 
comitants of  thought,  which  are  more  or  less  acces- 
sible to  us,  and  a  knowledge  of  which  may,  in  future, 
help  us  to  exercise  the  same  kind  of  control  over 
the  world  of  thought  as  we  already  possess  in  respect 
to  the  material  world;  whereas,  the  alternative,  or 
spiritualistic  terminology  is  utterly  barren,  and  leads 
to  nothing  but  obscurity  and  confusion  of  ideas. 
Thus,  there  can  be  little  doubt  that  the  further  science 
advances,  the  more  extensively  and  consistently  will 
all  the  phenomena  of  nature  be  represented  by  ma- 
terialistic formulae  and  symbols.  But  the  man  of 
science,  w^ho,  forgetting  the  limits  of  philosophical 
inquiry,  slides  from  these  formulae  and  symbols  into 
what  is  commonly  understood  by  materialism,  seems 
to  me  to  place  himself  on  a  level  with  the  mathema- 
tician, who  should  mistake  the  x's  and  y^s,  with 
which  he  works  his  problems,  for  real  entities — and 
with  this  further  disadvantage,  as  compared  with  the 
mathematician,  that  the  blunders  of  the  latter  are  of 
no  practical  consequence,  while  the  errors  of  sys- 
tematic materialism  may  paralyze  the  energies  and 
destroy  the  beauty  of  a  life." 

These  are  the  views  of  the  "physicists,"  so-called, 
a  school  represented  by  Prof.  Huxley,  Prof.  Owen, 
Herbert  Spencer,  Mr.  Grove,  Prof.  Tyndall,  and 
others.  Prof.  Owen,  in  the  last  pages  of  vol.  iii  of 
"  The  Anatomy  of  the  Vertebrates,"  declares  him- 


106  THE   CELL   DOCTRINE. 

self  the  champion  of  spontaneous  generation,  and  he 
maintains,  also,  that  the  formation  of  living  beings 
out  of  inanimate  matter  by  the  conversion  of  physi- 
cal and  chemical  into  vital  modes  of  force,  is  a  mat- 
ter of  daily  and  hourly  occurrence.  Mr.  Grove  says 
that  "  in  a  voltaic  battery  and  its  effects  we  have 
the  nearest  approach  man  has  made  to  an  experi- 
mental organism,"  and  that  in  the  human  body  we 
have  chemical  action,  electricity,  magnetism,  heat, 
light,  motion,  and  possibly  other  forces  "contribut- 
ing, in  the  most  complex  manner,  to  sustain  that 
result  of  combined  action  we  call  life.'* 

We  trust  it  is  not  necessary  to  state  that  it  has 
been  attempted  to  state  these  views,  not  in  a  spirit  of 
criticism,  but  fairly  and  justly;  our  object  in  this 
connection  being  simply  to  exhibit  the  present  state 
of  the  subject  as  viewed  from  all  standpoints.  And 
in  the  case  of  this  class  of  eminent  observers  we  have 
based  our  account  almost  entirely  on  what  we  believe 
the  latest  exposition  of  the  subject,  viz..  Prof.  Hux- 
ley's lecture,  while  we  have  included,  also,  such 
quotations  of  pregnant  sentences  of  other  observers 
of  the  same  class,  as  seemed  essential  to  complete- 
ness and  consistent  with  brevity. 

THE   author's   VIEWS. 

As  the  result  of  a  careful  comparison  of  the  views 
of  other  observers,  and  of  personal  observation,  ex- 
tending over  a  period  of  several  years,  chiefly  in 
the  direction  of  human  physiology  and  pathology, 
the  author  has  been  led  to  adopt  views,  which,  in 
the  main,  correspond  with  those  of  Dr.  Beale.    There 


THE   CELL   DOCTRINE.  107 

are,  however,  a  few  points  of  difference;  some,  per- 
haps, purely  in  mode  of  expression,  but  others  as  to 
matter  of  fact,  which  would  seem  to  be  appropriately 
here  recorded.  And,  in  order  to  give  completeness 
to  any  expression  of  such  views,  he  has  thought  best 
to  state  them  connectedly,  though  briefly. 

The  author  believes  the  ultimate  physical  element 
of  organization,  to  be  what  is  commonly  called  the 
"  cell,"  or  "  elementary  part,"  and  that  it  is  com- 
posed of  matter  in  two  states.  The  one,  central  in 
its  situation,  to  which  Dr.  Beale  has  most  appropri- 
ately given  the  name  "germinal  matter;"  the  other, 
for  the  most  part  peripheral  in  its  situation,  which 
the  same  observer  has  called  "  formed  matter."  The 
former,  which  is  the  "  sarcode  "  of  Dujardin,  the 
"protoplasm"  of  Max  Schultze,  is  that  upon  which 
the  origin  and  existence  of  the  cell  depends.  It  is  de- 
rived by  division,  budding  or  proliferation  from  pre- 
viously existing  matter  of  the  same  kind,  and  it  alone 
has  the  power  of  growing  by  converting  nutritious 
matter  or  "pabulum,"  derived  from  the  blood  or 
other  sources,  into  material  like  itself.  Without 
germinal  matter  textures  cannot  be  reproduced  or 
continued. 

In  appearance,  germinal  matter  is  often  structure- 
less, especially  as  constituting  the  living  moving 
matter  of  the  protozoa  or  lowest  animals  of  the 
rhizopod  type,  as  the  amoeba.  Yet  it  is  not  always 
structureless,  but  often  granular  in  its  appearance, 
and  as  constituting  the  mass  of  rapidly  growing 
cells  in  health  and  disease,  in  the  higher  animals, 
is  indeed  usually  granular,  as  is  evident  from  the 


108  THE   CELL   DOCTRINE. 

study  of  pus,  or  mucous,  or  white  blood  corpuscles, 
or  the  cells  of  a  rapidly  growing  morbid  growth.  In- 
deed it  seems  like  sacrificing  observation  to  theory, 
to  say  that  germinal  matter  is  always  structure- 
less. For  let  us  take  the  white  blood  corpuscle  or 
pus  corpuscle,  acknowledged  to  be  pure  germinal 
matter,  and  always  described  as  granular  in  its  struc- 
ture; either  the  germinal  matter  here  is  granular, 
or  the  granules  are  particles  of  formed  material  or 
extraneous  matter  suspended  in  the  formless  sub- 
stance, just  as  granular  matter  from  without  be- 
comes entangled  in  the  formless  matter  of  the  am- 
oeba. But,  such  a  view  as  the  latter,  would  be 
incompatible  both  with  the  behavior  of  growing 
germinal  matter,  and  the  reaction  by  which  it  is 
known;  for  we  note,  on  the  one  hand,  that  when 
germinal  matter  grows  rapidly,  these  granules  are 
the  elements  which  increase  most  abundantly;  and 
again,  that  these  are  the  portions  most  deeply  stained 
by  ammoniacal  solutions  of  carmine  or  aqueous  so- 
lutions of  red  aniline.  Especially  must  this  be  the 
case  if  the  so-called  nuclei  of  these  bodies,  which 
appear  after  the  addition  of  water  and  acetic  acid, 
are  simple  aggregations  of  the  granular  matter,  as  is 
contended  by  Dr.  Beale.  We  deem  it  incorrect, 
therefore,  to  describe  germinal  matter  as  in  all  in- 
stances structureless,  and  prefer,  with  Robin,  to  de- 
scribe it  as  sometimes  granular.  Indeed,  if  we  mis- 
take not.  Dr.  Beale  in  his  earlier  descriptions  also 
characterized  it  as  granular.* 

*  Beale's  Archives  of  Medicine,  vol.  ii,  p.  189. 


THE   CELL   DOCTRINE.  109 

A  circumscribed  round  or  oval  portion  of  germinal 
matter  within  the  cell  is  usually  termed  the  nucleus, 
which  may  be  surrounded  by  formed  material  as  in 
the  superficial  epithelial  cell,  or  by  other  germinal 
matter  as  in  the  white  blood  corpuscle. 

In  the  nutrition  of  the  cell,  the  pabulum  comes 
to  it  from  the  periphery;  being  strained  through  the 
formed  material,  and  the  new  germinal  matter  takes 
its  place  in  or  near  the  centre  of  the  original  mass, 
constituting  a  new  centre  of  germinal  matter,  which 
may  be  the  nucleus,  if  no  other  circumscribed  centre 
be  present,  or  the  nucleolus  if  it  be  deposited  within 
such  a  centre.  Other  new  centres  may  again  take 
position  within  these,  and  assume  the  relation  ot 
nucleolus  to  the  original  nucleolus,  which  now  be- 
comes the  nucleus,  an  older  centre  of  germinal  mat- 
ter; while  the  original  nucleus  has  probably  been 
converted  into  the  second  constituent  of  the  cell,  the 
formed  material. 

Germinal  matter  when  free  and  living,  exhibits  a 
power  of  movement,  both  in  portions  of  its  substance, 
producing  changes  in  shape,  and  in  its  entire  mass, 
resulting  in  changes  of  position.  The  former,  and 
probably,  also,  the  latter,  may  have  for  their  object 
the  obtaining  of  pabulum,  as  is  seen  in  the  amoeba, 
when  it  embraces  by  its  protrusions,  a  particle  of  nu- 
tritive matter.  These  movements  are  less  decided  in 
the  cells  of  the  higher  animals,  yet  they  are  of  con- 
stant occurrence,  as  in  pus  and  white  corpuscles,  and 
when  thus  occurring  they  are  spoken  of  as  "  amoeboid 
movements."  Allied  or  identical  with  this  second 
class  of  movements,  are  those  of  undoubted  occur- 

10 


110  THE   CELL   DOCTRINE. 

rence,  in  which  white  blood  corpuscles  have  been 
noted  by  Addison,*  Waller,t  and  Cohnheim,J  mi- 
grating from  the  bloodvessels,  and  constituting  one 
method  of  origin  of  pus. 

Formed  Material,  or  Non- Germinal  Matter,— As  the 
result  of  influences,  the  exact  nature  of  which  is  not 
known,  though  some  of  them  may  partake  of  the 
character  of  oxidations,  the  germinal  matter  is  con- 
verted into  the  second  constituent  of  the  cell,  formed 
material  This  formed  material,  peripheral,  for  the 
most  part  in  its  situation,  and  constituting  the  cell 
wall,  when  present,  is  without  the  property  of  ger- 
minating, or  multiplying  itself,  or  even  maintaining 
itself.  Yet  it  is  exceedingly  important,  and  as  essen- 
tial indeed  to  the  functions  of  the  economy,  as  the 
germinal  matter.  It  is,  in  fact,  the  portion  of  the 
cell  in  which  alone  function  resides,  since  it  is  to  the 
formed  material  of  the  muscle-cell  that  we  owe  the 
property  of  contractility,  to  the  formed  material  of 
the  nervous  element  that  we  are  indebted  for  neu- 
rility,  and  to  the  formed  matter  of  the  epithelial  cell 
that  we  owe  its  protective  qualities;  while  the  se- 
cretion of  all  glands,  whether  they  subserve  ulterior 
purposes  or  not,  is  the  formed  material  of  the  re- 
spective gland-cells.  Hence,  we  would  not  in  every 
instance  speak  of  the  formed  material  as  dead,  where 
it  is  the  seat  of  so  many  important  vital  endowments, 

*  Addison,  Physiological  Eesearches.     London  :  1841. 

f  "Waller,  London,  Dublin  and  Edinburgh  Philosophical  Maga- 
zine, vol.  xxix,  p.  271,  1846. 

X  Cohnheim,  Ueber  EntzUndung  und  Eiterung,  Virch.  Arch. 
Bd.  xl,  p.  48. 


THE   CELL   DOCTRINE.  Ill 

as  in  muscle  and  nerve.  In  some  situations,  it  is 
indeed  lifeless,  as  when  it  becomes  the  secretion  of 
glands,  as  bile  and  milk,  or  the  peripheral  part  of 
epithelial  cells.  It  simply  is  devoid  of  a  power  of 
multiplying  or  growing  by  itself,  depending  for  its 
increase  upon  the  conversion  of  the  germinal  matter. 
Hence  w^e  have  been  inclined  to  suggest  the  term 
"non-germinal,"or"  non-germinating"  matter,  since 
this  is  the  only  attribute  common  to  all  formed  ma- 
terial. 

In  structure,  formed  material  or  non-germinal  mat- 
ter is  varied.  Thus,  it  is  typically  without  structure 
in  the  red  blood  disc;  again  it  exhibits  distinctive 
structure  in  the  striped  sarcous  matter  of  muscle, 
and  in  the  fibrous  intercellular  substance  of  white 
fibrous  tissue  or  fibro-cartilage. 

As  formed  material  is  produced  on  the  periphery 
of  germinal  matter,  previously  existing  formed  ma- 
terial is  pushed  outward,  so  that  the  oldest  formed 
material  is  that  most  remote  from  the  germinal  mat- 
ter, and  the  youngest  lies  immediately  adjacent  to  it. 

Intercellular  substance,  whether  of  cartilage  or  white 
fibrous  tissue,  is  formed  material,  resulting  from  the 
conversion  of  the  germinal  matter,  which  constitutes 
the  cartilage  corpuscle  on  the  one  hand,  or  the  con- 
nective tissue  corpuscle  on  the  other.  It  is  not  of 
the  nature  of  a  deposit  from  the  bloodvessels  which 
subsequently  becomes  difi*erentiated.  Young  carti- 
lage cells,  like  all  young  cells,  consist  of  almost  pure 
germinal  matter,  and  the  capsule  of  the  cartilage 
corpuscle  is  but  formed  material,  more  or  less  con- 
tinuous and  inseparable  from  the  intercellular  sub- 


112  THE   CELL   DOCTRINE. 

stance;  so  that  we  would,  with  Beale,  define  a  carti- 
lage cell,  or  elementary  part  of  cartilage  as  com- 
posed of  germinal  matter,  with  as  much  surrounding 
formed  material  as  extends  half  way  to  the  adjacent 
germinal  matter.  So  with  the  elementary  part  of 
connective  tissue,  muscle,  and  nerve. 

Oil  and  starch  are  also  formed  matter,  conveniently 
designated  by  Dr.  Beale  as  secondary  formed  matter, 
and  result,  also,  from  a  conversion  of  the  germinal 
matter. 

As  already  stated,  the  proportion  in  which  these 
two  constituents  are  present,  is  various.  Thus,  in 
the  amoeba,  in  the  white  blood  disc,  in  the  pus  and 
mucous  corpuscle,  we  have  almost  pure  germinal  mat- 
ter, with  a  scarcely  appreciable  ring  of  formed  mat- 
ter on  its  periphery;  while  in  the  old  epithelial  cell 
we  have  almost  pure  formed  material  with  a  mere 
point  of  germinal  matter,  constituting  the  nucleus 
near  its  centre;  and  in  the  red  blood  disc,  we  have 
pure  structureless  formed  matter,  yet  matter  of  which 
we  should  long  hesitate  to  speak  as  dead.  In  old 
tendon,  again,  the  proportion  of  formed  material  is 
large,  and  germinal  matter  small,  while  in  young 
tendon  the  reverse  proportion  exists. 

The  cell,  as  thus  constituted,  and  originating  only 
in  the  germinal  matter  of  a  previously  existing  cell, 
we  believe  to  be  the  stariing-point  of  all  life  action,  be  it 
healthy  or  morbid.  Out  of  this  cell,  all  tissues,  simple 
and  complex,  are  constructed. 

We  believe,  also,  that  the  proper  shaping,  arrange- 
ment, and  function  of  these  elementary  parts  is  not  a 
process  identical  or  analogous  to  crystallization,  tak- 


THE   CELL   DOCTRINE.  113 

ing  place  through  merely  physical  laws,  but  that  there 
is  a  presiding  agency  which  controls  such  arrange- 
ment to  a  definite  end.  It  matters  not  what  this  is 
called,  but  we  prefer  to  designate  it  at  present  by  the 
term  "  vital  force,"  or  "  vitality."  It  is  this  controlling 
agency  which  makes  all  so-called  vital  properties  es- 
sentially different  from  purely  physical  properties,  a 
difference  which,  though  it  be  denied  in  ivords, and  ex- 
plained away  by  reasoning,  has  the  most  decided  proof 
of  its  existence  in  the  acknowledgment  it  receives  in 
the  actions  of  men,  just  as  the  most  convincing  argu- 
ment in  favor  of  the  free  agency  of  the  human  mind  is 
seen  in  the  fact,  that  all  men  shape  their  actions  on 
the  supposition  of  such  a  freedom,  whatever  their 
pretended  belief  with  regard  to  it. 

That  there  is  something  in  this  force  or  power  over 
and  above  the  physical  forces  of  nature,  is  most 
strikingly  shown  in  the  power,  exhibited  through  its 
agency  by  germinal  matter,  of  multiplying  and  pro- 
ducing new  germinal  matter  out  of  pabulum  unlike 
itself.  For  although  a  crystal  may  result  from  the 
rearrangement  of  particles  of  a  salt  in  solution,  as 
sulphate  of  alumina,  to  an  unlimited  extent,  there  is 
no  possibility,  nor  would  any  physicist  contend  that 
it  could  produce  crystals,  of  its  own  composition,  out 
of  carbonate  of  soda.  Nor,  as  is  justly  contended  by 
Dr.  Beale,  should  the  cell  be  compared  to  a  machine, 
unless  that  machine  possess  a  power  of  producing 
new  machines  out  of  material  unlike  itself,  and  of 
endowing  them  with  a  similar  power. 

In  morbid 'processes,  also,  the  germinal  matter  is  the 
seat  of  activity,  being  abnormally  increased,  dimin- 

10* 


114  THE    CELL    DOCTRINE. 

ished,  or  perverted;  and  many  pathological  states  are 
rationally  explained  by  bearing  in  mind  the  proper- 
ties of  germinal  matter  and  the  very  minute  size  which 
the  living  particles  may  exhibit.  All  physical  diffi- 
culties in  the  way  of  the  passage  of  white  blood  cor- 
puscles through  the  w^alls  of  capillaries  are  removed, 
when  we  remember  that  the  smallest  living  particles 
by  the  rapid  growth  of  which  white  blood  discs  or  pus 
corpuscles  are  speedily  produced,  do  not  exceed  the 
ttjtjVtju  of  an  inch  in  diameter,  and  that  however  un- 
reasonable it  may  appear  for  a  body  ^q\q  of  an  inch 
in  diameter  to  migrate  through  continuous  capillary 
walls,  it  becomes  much  less  unreasonable  when  we 
thus  reduce  its  proportions.  The  observations  of 
Beale  would  also  seem  to  reconcile  the  discordant 
views  with  regard  to  the  so-called  exudations,  in 
which  on  the  one  hand  we  need  not  suppose  an  ex- 
cessive dislocation  of  structure  to  admit  the  passage 
of  large  cells,  and  on  the  other  are  not  compelled  to 
restrict  the  origin  of  those  cells  to  points  outside 
the  vessels.  We  have  already  expressed  that  the 
views  of  H.  Charlton  Bastian  and  Cornil,  w^ith  regard 
to  the  origin  of  tubercle  in  the  perivascular  sheaths 
of  vessels,  are  not  practically  different  from  those 
earlier  expressed  by  Beale  as  to  its  origin  in  the 
germinal  matter  of  the  walls  of  bloodvessels. 

It  will  be  noted  that  the  only  points  of  difference 
between  our  own  and  the  views  of  Dr.  Beale,  lie  in 
the  structure  of  the  germinal  matter,  and  the  use  of 
the  word  dead  to  characterize  formed  material.  In 
all  other  respects,  we  accept  the  theory  of  Beale, 
and  have  no  hesitation  in  saying  that  it  admits,  with- 


THE   CELL   DOCTRINE.  116 

out  distortion  of  its  own  principle  or  disregard  of 
actual  facts,  of  consistent  application  to  a  larger  num- 
ber of  processes  of  tissue-building  in  health  and  dis- 
ease, than  any  other  theory  proposed. 

In  conclusion,  then,  it  may  be  stated,  1st,  that 
the  "  cell,"  or  "  elementary  part,"  originating  only 
in  a  pre-existing  cell,  is  the  ultimate  morphological 
element  of  the  tissue  of  animals  and  plants. 

2d.  That  the  cell,  contrary  to  the  belief  of  the 
earlier  histologists,  and,  indeed,  many  later  observ- 
ers, is  rarely  vesicular  in  its  structure,  but  generally 
more  or  less  solid  throughout. 

3d.  That  the  cell  is  composed  of  "germinal"  or 
living  matter  which  is  central,  and  includes  "  nu- 
cleus," "endoplast,"  "protoplasm"  and  "sarcode;" 
and  of  " non-germinal,"  or  "  formed"  matter,  which 
is  peripheral,  and  corresponds  with  "cell  wall"  and 
"intercellular  substance." 

4th.  That  this  germinal  matter  of  the  cell  in  a 
part  or  all  of  its  substance,  may  assume  a  special 
morphological  state,  usually  round  or  oval,  com- 
monly known  as  the  "  nucleus"  of  the  cell,  which, 
when  present,  is  always  a  young  centre  of  germinal 
matter;  but  that  in  other  instances  both  animal  and 
vegetable  cells  may  be  complete  without  this  special 
form  of  germinal  matter  or  "  nucleus,"  as  in  the  non- 
nucleated  amoebae  and  protogenes  primordialis  of 
Haeckel,  the  non-nucleated  monads  of  Cienkowsky, 
and  in  the  leaf  of  Sphagnum,  in  such  Algae  as  Hy- 
drodictyon,  Vaucheria,  and  Caulerpa,  and  in  young 
germinating  ferns. 

5th.  That  in  consequence  of  these  facts,  it  cannot 


116  THE   CELL   DOCTRINE. 

be  said  that  in  the  nucleus  alone  resides  the  power  to 
reproduce  the  cell,  since  we  find  the  nucleus  not 
essential,  but  that  in  the  germinal  matter,  of  which 
after  all,  the  nucleus,  when  present,  is  but  a  part, 
resides  this  function. 

6th.  That  when  the  smaller  body  within  the  nu- 
cleus, usually  known  as  the  "  nucleolus,"  is  present, 
as  it  often  is  in  complete  cells,  it  is  simply  a  younger 
centre  of  germinal  matter  than  is  the  nucleus  itself, 
and  is  the  last  formed  portion  of  germinal  matter, 
instead  of  being  the  oldest  part  of  the  cell,  as  orig- 
inally taught  by  Schleiden  and  Schwann.  And 
thus,  according  to  the  latest  views,  the  whole  process 
is  reversed.  The  old  order  of  succession  being,  1st. 
The  "nucleolus;"  2d.  About  this  the  "nucleus;" 
and  finally  about  this  the  "cell  wall,"  which  em- 
braces the  cell  contents.  Now,  however,  what  con- 
stitutes the  "cell  wall"  when  present,  is  the  oldest 
part  of  the  cell;  next  in  age  are  the  so-called  "cell 
contents,"  whether  germinal  matter  or  not;  next  the 
"  nucleus,"  and  last  and  youngest  the  "  nucleolus." 

7th.  That  the  formed  material  constituting  the 
cell  wall  and  intercellular  substance  may  be  some- 
thing chemically  different  from  the  germinal  matter, 
or  protoplasm  whence  it  was  converted,  as  the  secre- 
tions of  gland-cells,  or  may  be  a  simple  condensation 
of  the  exterior  of  the  cell,  as  iu  the  red  blood  disc. 

8th.  That  the  so-called  "  free  nuclei,"  so  often  re- 
ferred to  by  pathologists  in  their  descriptions  of  mi- 
nute structures,  are  simply  masses  of  germinal  matter, 
smaller  than  those  to  which  the  name  cell  is  usually 
given,  which,  if  time  be  permitted,  will  pass  into 


THE   CELL   DOCTRINE.  117 

perfect  cells  by  the  usual  production  of  formed  mat- 
ter on  their  periphery;  that  they  do  not  originate 
spontaneously,  but  from  previously  existing  germi- 
nal matter.  So,  too,  "  granules,"  if  they  be  com- 
posed of  germinal  matter,  present  the  same  attributes 
and  endowments,  arising  from  previously  existing 
germinal  matter,  capable  of  growing,  multiplying, 
and  assuming  all  the  characters  of  fully  formed  cells, 
but  never  originating  spontaneously.  Granules 
otherwise  composed  are  histolytic  (ktzo^,  a  tissue,  Xuatq, 
a  breaking),  and  7iot  histogenetic  (iffroq,  a  tissue,  y£V£(Tt<:, 
creation), — that  is,  they  result  from  the  breaking 
down  of  tissue  rather  than  go  to  building  it  up. 


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14 


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